You are on page 1of 187

B GIO DC V O TO

TRNG I HC NNG NGHIP I


PGS.TS. TRN C VIN - TS. NGUYN THANH LM
Ch bin: PGS.TS. trn c vin
gio trnh
sinh thi hc ng rung
NH XUT BN NNG NGHIP
H NI - 2006
1
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
LI NI U
ron
i
s p
hc lin h
T
g cc h sinh thi lc a th h sinh thi ng rung l ni c bin
t nhin su sc nht k t khi c loi ngi n nay. Trong lch
ht trin ca sinh thi hc, b mn c pht trin u tin l sinh
thi vi rng, sau vi ng c, ao h, sau cng mi xy dng c
sinh thi hc lin quan vi ng rung. Loi ngi bt u lm rung vo cui
thi i c (trc cng nguyn khong 7.000 nm), so vi lch s lu di
mt triu nm ca loi ngi th phi ni l kh gn. Trong qu trnh pht trin
nng nghip, tr tu loi ngi khng dng li vic iu khin mi trng sng
cho cy trng, m cn tin ln iu khin di truyn ca thc vt.
S pht trin ca nng nghip hin i t ra nhiu vn cn gii quyt.
Khuynh hng tng vic u t, thc cht l s u t nng lng ho thch,
thay th dn cho cc ngun li t nhin lm mi trng sng b hy hoi. Do
y, cn phi pht trin mt nn nng nghip da nhiu hn vo vic khai thc
hp l cc ngun li t nhin ca h sinh thi v bo v mi trng sng. l
nhim v s mt ca sinh thi hc nng nghip - c s ca vic b tr c cu cy
trng v vt nui hp l cc vng sn xut nng nghip.
S pht trin kinh t - x hi t ra yu cu ngnh nng nghip phi phn u
tng nng sut cy trng hn na. Rung cy trng nng sut cao l mt h
sinh thi hot ng hi ho, t c s cn i gia cc yu t cu thnh n. Do
y, thc cht ca k thut tng nng sut cy trng l k thut iu khin s hot
ng ca h sinh thi ng rung nng sut cao.
Gio trnh "Sinh thi hc ng rung" bao gm 5 chng. Chng I cung
cp khi nim chung v h sinh thi ng rung. Chng II m t cu trc v chc
nng ca h sinh thi ng rung. Chng III m t s vn ng ca h sinh thi
ng rung lin quan n cc nhn t nh sng, t ai, cy trng, c di, tun
hon vt cht v phn bn. Chng IV gii thiu cc bin php iu khin h sinh
thi ng rung. Chng V gip cho ngi hc, c bit l sinh vin sau i hc
h thng ho cc khi kin thc hc v cc k thut hc h thng ca h sinh
thi ng rung.
gip hc tt mn ny, trong tng chng c phn u gii thiu ni dung,
mc ch v yu cu i vi sinh vin. Sau mi chng, c trnh by phn tm tt,
cu hi n tp v ti liu c thm. Phn cui ca gio trnh l danh mc ti liu
tham kho v phn t vng (Glossary) m t cc khi nim v cc nh ngha
quan trng c s dng trong gio trnh ny.
2
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Gio trnh ny tng hp cc kin thc c ca nhiu mn khoa hc lin
quan dnh cho sinh vin bc i hc v sau i hc chuyn ngnh trng trt. im
mu cht ca gio trnh ny l gip cho ngi hc phng php t duy tng hp
thng qua cc phng php tip cn h thng c nhiu nc trn th gii p
dng thay th cho ci nhn n l trc y.
i vi sinh vin cao hc hoc nghin cu sinh, y l ti liu tt gip nh
nghin cu la chn cc bin php nghin cu ph hp.
Vi thi lng c hn dnh cho sinh vin bc i hc (2 n v hc trnh), nn
sinh vin cn nm c nhng nguyn l c bn ca sinh thi hc p dng cho
chuyn ngnh trng trt. ng thi nhng cng thc v phng trnh ton hc
trong gio trnh ny ch l ti liu b sung cho khi kin thc chnh v khng c
trong ni dung thi ca sinh vin. Trong qu trnh ging dy, gio vin s hng
dn sinh vin lng ghp kin thc v sinh thi hc ng rung vi cc khi kin
thc ri rc cc mn hc chuyn ngnh v c s thng qua cc bui tho lun
nhm v seminar.
Chng ti hy vng ngi hc s thy c, ch c trong mi lin h sinh thi
hc gia h sinh thi thin nhin vi h sinh thi ng rung th mi c th t c
s sng to ra h thng sn xut nng nghip bn vng v t tm ra mt con
ng duy tr nng sut cao trong nng nghip.
Do hn ch v thi gian v trnh , chc chn cun gio trnh "Sinh thi hc
ng rung" cn nhiu khim khuyt, rt mong nhn c nhng kin ng gp
ca cc ng nghip v ng o bn c. Tc gi xin chn thnh cm n v sn
lng tip thu cc kin ni dung gio trnh cng hon thin hn.


CC TC GI











3
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

1
Chng I
NG RUNG V SINH THI HC NG RUNG
Ni dung
Bn cnh sinh thi hc nng nghip, sinh thi hc ng rung ng gp mt phn
khng nh vo s pht trin mt nn nng nghip bn vng. y l mt mn khoa hc
tng hp nghin cu v mi quan h gia cy trng vi cc thnh phn sinh vt khc
(con ngi, ng vt, vi sinh vt, nm v c di) thng qua cc dng trao i vt cht,
thng tin v nng lng trong mi trng nh sng, nhit , khng kh, nc, m,
t ai,... Trong sut qu trnh pht trin ca nng nghip, cc sinh vt ch o trn
ng rung c s thay i su sc (v thnh phn ging, nng sut, kiu hnh, cht
lng, kh nng chng chu, v.v...). ng thi, con ngi to ra cc vng phn b
c trng cho cc kiu ng rung khc nhau. Chnh v vy, cc h sinh thi ng rung
xut hin vi cc cu trc v chc nng c trng c th cho tng vng.
Cc ni dung c cp trong chng I:
Khi nim chung v sinh thi hc ng rung.
Qu trnh hnh thnh v pht trin ng rung.
Cu trc v chc nng ca h sinh thi ng rung.

nh. H canh tc nng ry tng hp ca ngi Ty Bc, Ha Bnh:
Rung bc thang thung lng, nng la v sn trn lng chng i
v rng ti sinh trn nh i
Mc tiu
Sau khi hc xong chng ny, sinh vin cn nm vng:
Khi nim, ni dung, i tng ca sinh thi hc ng rung.
S hnh thnh v pht trin ca ng rung.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

2
1. Khi nim chung v sinh thi hc ng rung
1.1 ngha v tc dng ca sinh thi hc ng rung
T trc n nay c nhiu nghin cu v mi trng i vi cy trng - i
tng ca sn xut nng nghip. Nhng phn nhiu l nhm vo nhng nh hng ca
iu kin mi trng ring bit nh th nhng, kh hu, c di... i vi cy trng; rt
t nhng nghin cu coi ng rung l mt h thng c cu thnh t loi ngi cho
n vi sinh vt.
H sinh thi ng rung c t ngang hng vi cc h sinh thi t nhin nh rng,
ng c, vc nc, lc a... Thut ng h sinh thi ng rung mi gn y mi c
c v tr r rng trong sinh thi hc ng dng. H sinh thi ng rung l mt h thng
vi qun th hoc cc qun th cy trng l trung tm tng tc cht ch vi mi trng
xung quanh bao gm nh sng, khng kh, nc, a hnh, t ai, c di, cn trng, vi
sinh vt, ng vt, v.v... (hnh 1.1). H sinh thi ng rung l mt trong nhng h sinh
thi trong sinh quyn, v vy v nguyn tc, phng php v cch nghin cu sinh thi
hc u thch hp vi n. Ngc li, nhng quy lut v phng php m sinh thi hc
ng rung tm ra cng c th vn dng cho cc lnh vc sinh thi hc khc.
So vi sinh thi hc nng nghip hay sinh thi hc cy trng m ni dung ch yu
l a l hc sinh thi cy trng th sinh thi hc ng rung c c im l c tin
hnh nghin cu trong h sinh thi mt cch tng hp v ng hn.















Lin h ln nhau gia
cc h sinh thi trong
h sinh thi lc a


Qu trnh sn xut ca
qun th cy trng trong
h sinh thi ng rung


Cy
trng
H sinh thi
ng rung
Nghin cu v nu r
h sinh thi trong mi trng
do ngi iu khin
Lp t (cm)

Phn tch tng hp
ng thi h sinh thi
ng rung (lin h ln
nhau ca ton b cc
thnh phn h sinh thi)
Hnh 1.1. Phm vi nghin cu ca sinh thi hc ng rung
Mt khc, ni v s cn bng ton b h thng, trnh iu khin k thut nng
nghip hin nay i vi h sinh thi ng rung cn cch xa mi c nh mong mun.
Th d, khi chng ta tm cch c ch mt loi c di no pht trin th c th dn n
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

3
lm mt loi c di khc pht trin; iu khin di truyn vn l tng sn lng vt
cht kh ca cy trng, nhng kt qu c khi ngc li l gip lm tng sinh khi ca
loi c hi; hay cy su qu mc li lm gim kh nng sn xut ca ng rung do
lm bc phn. Tm li, c th thy tc dng ca sinh thi hc ng rung l thng qua
vic gii thch h sinh thi ng rung m tm ra quy lut hay phng php nghin cu
h sinh thi khc. Nghin cu mt cch ng v tng hp theo cch tip cn h thng
trong mi lin h gia h sinh thi ng rung vi cc h sinh thi khc xung quanh n
tm ra con ng iu chnh nng cao nng sut sn xut v ch o k thut sn xut
c th (Altieri, 2002, Phm Ch Thnh v ctv, 1996, Trn c Vin, 1998).
1.2. Cc hng nghin cu chnh ca sinh thi hc ng rung
Sinh thi hc ng rung tp trung nghin cu theo ba hng: (1) Qu trnh sn
xut ca qun th cy trng trong h sinh thi ng rung; (2) Gii thch tng hp ng
h sinh thi ng rung (lin h ln nhau ca ton b cc phn hp thnh); (3) Quan
h ln nhau gia h sinh thi ng rung vi cc h sinh thi khc xung quanh n
(Trn c Vin, 1998).


















Qun th cy trng



Quang hp Vn chuyn Chn

Chuyn ho
nng lng
Phn phi
nng lng
Ct gi
nng lng
Nc
CO
2
C
di
Nng
lng
mt tri
Nm
bnh
Su
hi
Nm
bnh
Su
hi
Nc
Cht
dinh
dng
Hnh 2.1. Quan h ca chuyn ho nng lng v cc phn hp thnh
trong h sinh thi ng rung ly qun th cy trng lm trung tm
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Ni dung nghin cu ca sinh thi ng rung ht sc rng, bao gm ni dung
nghin cu t mc c th n mc gia cc h sinh thi. Cng nh cc h sinh thi
khc, qu trnh c bn ca h sinh thi ng rung l s chuyn ho nng lng v tun
hon vt cht ca h sinh thi m ht nhn l sinh vt. Chi tit s cp phn cui
chng ny, y ch nu mt th d, trong h sinh thi ng rung biu th bng hnh
v, chuyn ho nng lng ly qun th cy trng lm trung tm, nng lng mt tri
chiu trn ng rung di nh hng ca cc thnh phn hp thnh khc nhau ca h
sinh thi, thng qua qun th cy trng m tin hnh trao i, c nh, phn phi v ct
gi (hnh 2.1). Do , tu theo vic nu r nhng nhn t v qu trnh, li lm r thm
s din bin v qu trnh thch ng ca c h sinh thi, gii thch kt cu, chc nng v
ng thi ca n, ng thi p dng phng php ca k thut hc h thng phn tch
c h sinh thi, mi c th hiu h sinh thi ng rung mt cch ton din hn.
2. Qu trnh hnh thnh v pht trin ng rung
2.1 Bt u ca nng nghip l s hnh thnh v pht trin ng rung
Theo hiu bit hin nay, loi ngi c xa nht (Australopithecus) sinh ra vo thi
k bng h th nht (trc cng nguyn khong 1 triu nm), qua thi k bng h th t
(trc cng nguyn khong 25.000 nm) cho n cui thi i c (trc cng
nguyn khong 7.000 nm), mi tin ho thnh loi ngi ngy nay (Homosapiense),
thi gian khong 1 triu nm. Trong thi gian , loi ngi sinh sng da vo sn bt
v hi lm, sau mi bc vo thi i chn nui v lm rung. Do s khc nhau
gia cc khu vc, vic phn chia ra cc thi i sn bt, chn nui, lm rung cn ph
thuc vo iu kin thun li v kh khn ca tng vng. Nhng d th no, lm rung
vn l khuynh hng lm cho loi ngi nh c thnh b lc, ng thi cng thc
y pht trin nhanh k thut sn xut nng nghip v pht trin vn ho-kinh t-x hi.
Bui u ca k thut ngh nng, theo truyn thuyt, l c gi ra t ht cc vi
trong t ny mm v sinh trng. Nng c c xa nht l cy gy v ru o t,
iu ny cng ni r k thut ngh nng u tin l bt u t vic o lt t (Peake,
1928; Kumadai, 1970). Cy lt t l cch iu khin do con ngi bt u tc dng vo
h sinh thi t nhin nh rng, ng c, bi sng..., kt qu l sinh ra ng rung
chung quanh nh ca nhng ngi nh c. Nng ry t l hnh thi u tin ca
tng mnh rung, tc l t ry, chc l b ht, t n thu hoch khng chm sc g
c, qua nhiu nm t ngho i th b ho chuyn n ni khc, li tin hnh kiu nng
nghip bc lt t nh vy. Vit Nam n thi cc vua Hng c nn nng nghip
rung nc, th hin qua cc truyn thuyt nh cu chuyn v Sn Tinh, Thy Tinh.
Trn mt t ca h sinh thi nng ry c tro l ngun phn v c phong ph.
Ngoi ra, la c th t cht nm bnh, cn trng, vi sinh vt v c di, c li trong mt
thi gian ngn. t nng ry trong thi gian b ho, c ngh v hi phc dn. Nn
nng nghip nguyn thu ny n nay vn cn tn ti nhiu vng nhit i.
4
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Sau ny cng vi s pht trin v thay i ch s hu t ai, duy tr mu
m ca t, ngi ta p dng phng thc cho t ngh. ng thi nng cao
mu m ca t v mc s dng t, ngi ta tin hnh lun canh. ng rung c
chia thnh nhiu mnh, c t ngh, t gieo v cc xun, t gieo v cc ng. Do k
thut cy ba pht trin, t ngh khng c ngha l khng qun l, m vn cy ba
tr c di v ci thin iu kin thng thong cho t, ngha l p dng nhiu cch tch
cc khi phc mu m ca t. Hn na, do tin b ca k thut lun canh, ngoi
cy cc ra, cn c thm nhiu loi cy trng khc, h sinh thi ng rung ngy cng
thm phc tp (Grass, 1925; Orwin, 1949).
Cng vi nng ry v ch canh tc rung nc vng t cao, vng t thp
cng pht trin ng rung. Cc dn tc nh c vng ven h, sng, m ly, qua giai
on nh bt c, lm hi qu, cng hc cch lm rung. Nn vn ho phng ng
c i pht trin bn cc dng sng ln nh Tigr, frat, Nil; nn vn ho dn tc
Hn bt ngun t lu vc sng Hong H; nn vn minh la nc ca ngi Vit
vng ng bng sng Hng ..., u ly c s l vn minh nng nghip.
Theo ti liu kho c, s pht trin ca ng rung l t vng t cao xung vng
t thp (Furusima, 1947; Marukufuchi, 1968). Nng nghip t thp cn gi l nng
nghip t ngp nc, mt mt c nhng cht lng ng ph sa do dng sng chuyn
ti hnh thnh ng rung mu m, mt khc k thut iu khin ngp lt ca nc
sng c pht trin khng ngng theo s pht trin ca th ch chnh tr, x hi, nh
Vit Nam hnh thnh h thng iu v h thng dn thu nhp in. V th, h
sinh thi ng rung ca nng nghip t thp, thng qua cc bin php tr thu v dn
ti, ly vic khng ch nc lm trung tm, duy tr cn bng sinh thi ca c h thng
(Morimoto, 1941; Marukufuchi, 1968).
2.2. S phn b ca ng rung
Nc l mt trong nhng nhn t hn ch s tn ti hay khng tn ti ca ng
rung, do , i chiu s phn b ma ca lc a cho thy: vng c lng ma nh
hn 250 mm hu nh khng c din tch ng rung. Ngay la m, mt cy trng chim
din tch ng rung rng nht th gii vi tng s 257 khu vc trng th 83% s khu
vc l nm trong vng c lng ma 500 - 1000 mm/nm.
S phn b ca ng rung, ni mt cch khi qut l do nc, nhit , a hnh
v v hn ch. i vi cc loi cy trng ch yu th b cc iu kin mi trng nht
nh lin h vi ging cy hoc quan h t hp ca mt s iu kin no hn ch.
tng khu vc cc b thng b nh hng ca su bnh. V th, v mt quy hoch t
thch hp vi cy trng, vic nghin cu a l sinh thi cy trng l mt trong nhng
lnh vc quan trng ca sinh thi hc ng rung. Vit Nam c hai va la chnh l
ng bng sng Hng v ng bng sng Cu Long, l hai vng a hnh bng
phng nht so vi cc vng khc trong c nc.
5
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Bng 1.1. Din tch trng la ti 4 vng sn xut la chnh ca c nc
theo a hnh tng i
a hnh tng i
Vng
Din tch
(1000 ha)
Cao Vn cao Vn Vn thp Trng
ng bng sng Cu Long 2082,7 87,9 813 1073,3 108,5
ng bng sng Hng 667,3 6,6 64,9 227,4 291,6 76,8
Duyn hi Bc Trung B 395,8 31,6 91,1 177,8 66,9 28,4
Duyn hi Nam Trung B 279,9 15,1 63,7 55,6 117,9 27,6
Tng cng 3425,7 53,3 307,6 1273,8 1549,7 241,3
Ngun: Vin Quy hoch v Thit k nng nghip, 2003
Theo s liu trong bng 1.1, din tch la nc tp trung ch yu hai vng ng
bng chnh l ng bng sng Cu Long v ng bng sng Hng. ng thi, do iu
kin a hnh nn din tch la nc vng Bc Trung B v Nam Trung B c din
tch nh hn rt nhiu, ch bng mt na din tch la ng bng sng Hng v 1/7-1/8
din tch la ng bng sng Cu Long. mi vng, la nc li c chia ra theo
a hnh tng i nh cao, vn cao, vn, vn thp v trng (Vin Quy hoch v Thit
k nng nghip, 2003).
2.3. S hp thnh ca h sinh thi ng rung
im khc nhau ch yu v thnh phn hp thnh ca h sinh thi ng rung so
vi h sinh thi khc l qun th cy trng mang tc dng ch o do con ngi iu
khin mt cch y ; ngi v gia sc cng l thnh phn hp thnh ca h sinh thi.
Ngoi ra, cn c mt s bin php iu khin ca con ngi c nh hng su sc n
s hp thnh ca h sinh thi ng rung nh bin php lm t, bn phn, phng tr
su, bnh, c di, ph t, ti nc v iu khin di truyn chn ging (Trn c
Vin, 1998; Phm Ch Thnh v ctv, 1996).
S hp thnh ca h sinh thi ng rung nh trong hnh 1.1, l qun th cu thnh
bi cy trng n nht (ging) v cng vi qun th cy trng ch o l qun th cy
trng khc, qun th c di, su hi, ng vt ..., to thnh qun x sinh vt ca h sinh
thi ng rung. Gia cc thnh phn hp thnh ny tn ti mi quan h qua li phc
tp (s ni k phn sau), y ch nu mt th d. Nh trong bng 2.1, trong qu trnh
chuyn bin t h sinh thi t nhin (ng c) sang h sinh thi ng rung (t la m),
qun th cn trng, thnh phn hp thnh ca h, b nh hng r rt. T ng c bin
thnh t la m, nm th nht, s loi cn trng u gim i, nhng tng s c th u
6
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

tng ln. Nu tnh theo loi, nh phn di ca bng 2.1, s tng gim s c th ca cc
loi khc nhau, c mt s loi c tc dng c ch i vi loi khc, s c th ca chng
tng ln, kt qu chung l lm cho s c th tng ln.
Bng 2.1. nh hng ca s chuyn bin t h sinh thi t nhin sang h sinh thi
ng rung i vi thnh phn qun th cn trng (WATT, 1968)
Tn cn trng ng c t la m
Tng gim
(ln)
S loi
Homoptera
Heteroptera
Coleoptera
Hymenoptera
Loi khc
Cng cc loi
Tng s c th/m
2

35
38
93
37
137
340
199

12
19
39
18
54
142
351

- 2,9
- 2,0
- 2,4
- 2,1
- 2,5
- 2,4
+ 1,8
S c th ca cc loi khc nhau/m
2
Caeculus dubius
Sminthurus viridis
Ectobius duskei
Leptothorax nassonovi
Haptothrips tritia
Phyllotreta viitula
Hadena sordida

2,23
7,65
2,83
16,48
1,07
0,05
0,09

0,06
0,19
0,13
0,03
300,40
1,03
2,25

- 37,2
- 40,2
- 94,3
- 550,0
+ 280,8
- 20,6
+ 25,0
H sinh thi ng rung l h thng phc tp cu thnh bi cc thnh phn v c
v cc thnh phn sinh vt, nn phi phn tch ng thi mi quan h ln nhau gia cc
thnh phn hp thnh mi c th gip k thut trng trt tng hp nng cao sn
lng qun th cy trng.




7
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

TM TT
H sinh thi ng rung l mt trong nhng h sinh thi trong sinh quyn, v vy,
phng php v cch nghin cu sinh thi hc v nguyn tc l u thch hp vi
n. Ngc li, nhng quy lut v phng php m sinh thi hc ng rung
tm ra v nguyn tc cng c th vn dng cho cc lnh vc sinh thi hc khc.
Sinh thi hc ng rung hin ang tp trung nghin cu theo ba hng: (1)
Qu trnh sn xut ca qun th cy trng trong h sinh thi ng rung; (2)
Gii thch tng hp ng h sinh thi ng rung (lin h ln nhau ca ton
b cc phn hp thnh); (3) Quan h ln nhau gia h sinh thi ng rung
vi cc h sinh thi khc xung quanh n.
Lch s pht trin ng rung tri qua mt thi gian di. Lm rung vn l
khuynh hng lm cho loi ngi an c, lc nghip, ng thi cng thc y pht
trin nhanh k thut sn xut nng nghip v pht trin vn ho- kinh t- x hi.
S phn b ca ng rung, ni mt cch khi qut l do nc, nhit , a
hnh, t ai v kh hu hn ch. i vi cc loi cy trng ch yu th b cc
iu kin mi trng nht nh lin h vi ging cy hoc quan h t hp
ca mt s iu kin no hn ch nh kinh t - x hi ca tng vng.
tng khu vc cc b thng b nh hng ca su bnh. V th, v mt quy
hoch t thch hp vi cy trng, vic nghin cu a l sinh thi cy trng l
mt trong nhng lnh vc quan trng ca sinh thi hc ng rung.
im khc nhau ch yu v thnh phn hp thnh ca h sinh thi ng rung
so vi h sinh thi khc l qun th cy trng mang tc dng ch o do con
ngi iu khin mt cch ton din; Thnh phn loi sinh vt trn ng
rung c s thay i su sc nu con ngi thay th cy trng ch o. Nhng
tc ng ny c nhng nh hng khc nhau n li ch con ngi.
CU HI N TP
1. c trng c bn ca h sinh thi ng rung l g?
2. Phm vi nghin cu ca h sinh thi ng rung?
3. Ti sao hin nay sinh thi hc ng rung tp trung vo 3 hng nghin cu chnh?
4. Ti sao cc h sinh thi ng rung li khc nhau theo cao v khc nhau theo vng?
5. S khc bit cn bn v thnh phn loi gia h sinh thi ng rung v h sinh
thi t nhin?
TI LIU C THM
1. Phm Ch Thnh, Phm Tin Dng, o Chu Thu, Trn c Vin. H thng
nng nghip. NXB Nng nghip. H Ni - 1996.
2. Trn c Vin. Sinh thi hc nng nghip. NXB Gio dc. H Ni - 1998.
8
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Chng II
CU TRC V CHC NNG CA H SINH THI NG RUNG
Ni dung
Trong lch s nghin cu h sinh thi rng v ng c t nhin, vic nghin cu
cu trc qun x thc vt c pht trin tng i sm, iu c tc dng nht nh
i vi vic xy dng khi nim qun x thc vt. Khi nim cu trc qun x phi bao
gm: cc loi hp thnh v kiu sinh sng ca chng, s phn b khng gian ca chng,
s phn b v lng o bng i lng hay ch s no (nh mt , tn , trng
lng...) v nhng bin i ca chng theo thi gian...
H sinh thi ng rung, tr qun x c di ra, thng rt n gin, tc l qun th
cy trng ch do mt loi cu trc thnh. Mt khc, h sinh thi cy trng ly qun th
cy trng lm chnh cng vi cc thnh phn ph nh qun th c di, ng vt, qun
th vi sinh vt v mi trng vt l. V th, khi nu r cu trc v chc nng ca h
thng, khng ch gii hn cu trc ca qun th cy trng, cn phi lm sng t cu
trc qun th sinh vt khc, mi trng vt l v ng thi tc dng gia chng vi
nhau.
Cc ni dung sau y s c cp trong chng ny:
1. Cn bng lng nhit v cn bng nc ca ng rung
2. Mi trng t
3. Mi trng sinh vt
4. Cu trc ca qun th cy trng
5. Cu trc mi trng ca h sinh thi ng rung
6. Quang hp ca qun th cy trng
7. S sinh trng ca qun th cy trng
8. S cnh tranh trong h sinh thi ng rung
9. Nng sut ca h sinh thi ng rung
10. M hnh ha h sinh thi ng rung.
Mc tiu
Sau khi hc xong chng ny, sinh vin cn:
1. Hiu c cu trc v chc nng ca h sinh thi ng rung,
2. Hiu c mi trng t, mi trng sinh vt ca h sinh thi ng
rung,
3. Hiu c mi quan h gia c di v cy trng, s sinh trng ca qun
th cy trng trong h sinh thi ng rung.
1
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

2
Nng lng ca s vn ng suy cho cng u bt ngun t nng lng mt tri,
do nghin cu tc dng ca mi trng vt l v qun th cy trng i vi qu trnh
trao i nng lng mt tri tng khng kh gn mt t c th nu r cu trc mi
trng ca h sinh thi ng rung. Cu trc ca h sinh thi ng rung rt phc tp,
quyt nh cu trc v chc nng ca h thng. Th d, quang hp ca qun th cy
trng, cu trc ca qun th b mt tng l v phn b khng gian ca tng l quyt
nh. Nhng quang hp li hnh thnh l mi, lm thay i cu trc tng l v li nh
hng ti chc nng v cu trc ca h thng. Quan h ny c ngha l: khng c nh
lng cu trc ca h thng s khng th nu r mt cch nh lng chc nng ca h
thng. Xut pht t quan im i vi cu trc mi trng nh vy, di y s nu r
vn cn bng lng nhit v cn bng nc ca ng rung, vn biu hin nh
lng cu trc ca h thng v hm s ho chc nng ca h thng.
1. Cn bng lng nhit v cn bng nc ca ng rung
1.1. Cn bng lng nhit ca ng rung
Nghin cu s trao i nng lng mt tri trn ng rung, c bn nht l nghin
cu v cn bng bc x v cn bng lng nhit. Cn bng bc x l tng bc x nng
lng mt tri, khng kh v mt t, c ngha l nhit nng thun m mt t ng
rung thu c, cng gi l bc x thun. S bin i nng lng mt tri ch yu vi
hnh thc nhit, cho nn cng dng thut ng cn bng lng nhit lm t ng ngha
ca thut ng cn bng nng lng.
















+9
My
Khng
kh
+10
33
10 56
109
105 Khng
kh
+ 23
Khng
kh
Khng
kh
+ 10
Bc x mt tri
Bc x mt t
v khng kh
Bc hi v
ngng t
25 52 15 9
17 24 6
-66 -9 +100 -25
i lu v
truyn dn
+17 + 24 +6 -119 +105 -23 -10
100 n v = 0,485 cal/cm
2
/min
Hnh 1.2. Cn bng lng nhit ca mt t (Gates, 1962)
Trao i nhit
lng thun
+ 47 -23 -10 -14
Tim nhit Cm nhit Bc x
sng di
Bc x
tn x
Bc x thng
quang my
Bc x
trc tip
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

3
Bnh qun nm ca cn bng nhit lng ng rung:
Hnh 1.2 l tnh hnh phn phi li ca nng lng mt tri biu th bng tr s bnh
qun nm ca Bc bn cu. Ly bc x mt tri l 100, tr s ny tng ng vi
0,485 cal/cm
2
/pht, trong chiu trc tip xung mt t 33, khng kh hp th 49,
n mt t 24 tr thnh bc x trc tip; 52 n b mt my, t mt 25 phn x vo
trong khng gian v tr, 10 c my hp th, 17 thng qua my n mt t. Mt
khc, 15 n v ta mt trong khng kh, 9 n v to vo v tr, cn 6 n v n mt
t, cng vi nh sng thng qua my n mt t ni trn thnh bc x tn lon (tn
x). Kt qu l nng lng mt tri chiu vo tng trn khng kh ch c 47% n c
mt t, 34% phn x vo khng gian v tr.
T mt t chiu ra bc x nhit sng di 119, trong 10 i vo khng gian v
tr, s cn li c khng kh hp th. T khng kh li vi bc x sng di 105 n
mt t. Do , lm trao i lng nhit sng di trn vn b mt i 14 t mt t.
Do c 56 bc x sng di t khng kh chiu vo khng gian v tr, cho nn ton b
nhit nng m qu t mt vo v tr l 100, lm cho nhit ca ton th qu t
khng ln cao. Trong s 47 n mt t, c 23 l lng nhit bc hi ri tiu tan trong
khng kh. Loi lng nhit lu ng to ra v hp th khi nc bc hi v ngng t
gi l tim nhit. Cui cng cn li 10 c gi chuyn vn, thng qua i lu v
truyn dn, nm trong khng kh, loi nhit nng ny gi l hin nhit.
Hnh 2.2. Quang ph bc x mt tri (a). So snh ng cong quang ph
quang hp ca la m v cm gic nhn thy tng i (b)
1. Quang hp ca la m (Gates, 1962); 2. cm gic nhn thy tng i (Laisk, 1965)
2,8 3,2 2,4 2,0 1,6 1,2
Bc x n ca 6000
0
K
Bc x mt tri ngoi kh quyn
Bc x mt tri ca mt t
O
2
O
3
H
2
O
3,0
0,4 0,8
O
3
Bc sng




0,7 0,5 0,4
1
2
Bc sng
0,6
H
2
O
H
2
O
Hng
ngoi
H
2
O
Nhn thy
0
0
1,0
2,0
UV

a)


C

n
g

c

x









b)
2,0

C

n
g

n
g

i

2,0

2,0
2,0

2,0

0



Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Cng thc cn bng nhit lng ng rung:
Hnh thi ca nng lng mt tri tuy c bin i, nhng theo nh lut bo ton
nng lng th khng mt i. Vy cng thc cn bng lng nhit nh sau:
R + H + IE = 0 (1)
Trong : R: Bc x thun; H: Cm nhit; I: Tim nhit bc hi; E: Lng bc hi
trn n v din tch, n v thi gian.
Nu xt n nhng bin i trong thi gian tng i ngn, trn ng rung, th
cng thc trn c th i thnh:
R + H + IE + B + P = 0 (2)
Trong : B l nhit tn tr trong ng rung, dng vo s ln xung nhit t
v nhit thn thc vt; P l nhit tn tr ha nng ca quang hp.
Tr s ca chng rt nh so vi cc s hng khc, hu nh c th b qua.
Bc x thun: c th biu th bng cng thc sau y:
R = (1 - a) (Q + q) + S (3)
a l sut phn x ca ng rung
Q v q l bc x mt tri chia ra trc tip v tn lon
S l bc x hu hiu sng di, l tng ca bc x sng di t mt t ra
v t khng kh n.
Du ca cc s hng trong cc cng thc t (1) n (3) ly chiu chiu vo mt t
l dng, chiu phn x l m.
Bc x mt tri v bc x quang hp c:
V i th, bc x mt tri gm c bc x bng sng 0,2 - 4,0, gi l bc x sng
ngn, cng ca cc bc sng khc nhau nh hnh 2.2 (a) cho thy, ngoi kh
quyn th gn nh bng bc x t ngun 6000
0
K, cn trong khng kh th c hi
nc, oxi, ozon, bi hp thu, hnh thnh my khe lm. Trong , bng sng cho quang
hp c, nh hnh 2.2 (b) cho thy, gn bng vi phn nhn thy c: 0,38 - 0,71.
Bc x ca bng sng ny gi l bc x quang hp c. Hnh 2.2 cn cho bit, quang
ph c tc dng i vi quang hp v ng cong biu th cm gic ca mt ngi
i vi cc bc sng khc nhau r rng l khc nhau. Do , khi o quang hp, dng
lux biu th cng nh sng l khng chnh xc.
Bc x quang hp c i th tng ng vi mt na ca bc x mt tri.
Tooming v Guliaep (1967) cho rng, tr s tnh tch ngy v tr s tnh tch thng c
quan h nh sau:
Q = 0,42 Q + 0,60 Q (4)
Trong : Q l bc x quang hp c. T cho thy, t l bc x quang hp
c ca bc x tn x l cao hn.
4
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

5
Thng lng hin nhit v tim nhit:
B<0 B>0
H>0 H>0
B>0 B<0
H<0
H<0
nhit
Khng kh
t
cao







Mt t



Hnh 3.2. Bin i trong ngy v phn b nhit ng rung
Hnh 3.2 cho thy, bin i trong ngy theo chiu thng ng ca nhit khng
kh v nhit t mt t vo ban m l thp nht, bui tra tr nn cao nht. Bin
i trong ngy ca nhit t c th n su khong 30cm, su hn na hu
nh khng c bin i nhit. Nhng cn c vo ng cong bin i trong nm ca
nhit t th s bin i nhit c th n su 600 cm. Xem dc ca ng
cong trong hnh c th thy hin nhit ca mt t, hng truyn dn ca nhit truyn
dn trong t v mc ln nh ca nhit, nh mi tn trong hnh. Truyn dn nhit
thnh t l vi dc ca ng cong (
dz
DT
), dc cng ln th dn nhit cng nhiu.
dc nhit v thng lng hin nhit mt thi gian no c quan h nh sau:
H = C K
H
dz
DT
(5)
Trong : l mt khng kh; C l t nhit nh p ca khng kh;
K
H
l h s khuch tn x
dz
DT
t l bin i nhit theo cao ( dc thng ng).
Cng nh trn, tim nhit c th biu th nh sau:
IE = K
W
I
dz
dq
(6)
Trong : q l m tng i, K
W
l h s khuch tn dng xoy ca hi nc.
Hai cng thc (5) v (6) cho bit: h s khuch tn dng xoy cng ln, dc
thng ng ca nhit hay m cng ln, th thng lng hin nhit v tim nhit s
cng ln.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Lng bin i nhit tn tr:
Nhit tn tr trong ng rung (B) biu th s thay i lng nhit ca mt ct cu
thnh do r cy trng v t tnh t mt t tr xung, cng tc l thng lng nhit tn
tr. Trn mt t trong qun th cy trng, c th theo cng thc (2), tc l:
R
0
+ IE
0
+ H
0
+ B
0
= 0 (7)
Trong cng thc ny, B
0
biu th nhit truyn dn t mt t xung di t v gi
l nhit truyn dn trong t, c th biu th bng cng thc sau y:

0 z
dz
dTS
B0 =
(8)
Trong : l h s dn truyn nhit ca t (cal/cm
2
. sec.
0
C);

0 z
dz
dTS
=
biu th dc thng ng phn b nhit t ca mt t.
Tr s B
0
tin hnh phn tch 1 nm th gn bng khng.
T (2) n (7) ta c: Bp = B - B
0
(9)
Trong : Bp l lng bin i nhit tr trong thn cy trng v trong khng kh
ca qun th cy trng.
S phn b a l v cn bng lng nhit:
Nh kh hu hc Lin X Buuko dng phng php kh hu hc nghin cu s
phn b a l v cn bng lng nhit: lng nhit to ra do bc hi, lng trao i
nhit khuch tn. Nghin cu ch ra rng, v mt lng nhit ca c nm, nh hnh 4.2
(a) cho thy, bc x thun trn lc a v trn bin khc nhau r rt, ng ranh gii
biu th tr s khng lin tc. l do t l sut phn x ca mt bin nh hn so vi b
mt lc a cng v . Tr s ln nht ca bc x thun trn qu t c thy phn
Bc bin Arabi, khong 140kcal/cm
2
.nm. bin, ng thng tr thnh tng bng ko
di hng ng Ty; i v cao, tr s bc x thun h thp rt nhanh.
Trn lc a, tr s bc x thun ln nht c thy vng nhit i m, cng ch
c 100 kcal/cm
2
nm, rt nh so vi trn bin. Ngoi ra, nu so snh vng kh vi vng
m th tr s nh hn. l do sut phn x ca bc x sng ngn vng kh ln hn,
bc x hu hiu sng di cng ln ( nhit b mt cao, ngy rm t, m thp).
Hnh 4.2 (b) biu th nhit to ra do bc hi. Tr s ca lc a v bin khc bit
nhau r rt, thay i r rt ng ranh gii. y cng ging vi tnh hnh bc x
thun ni trn, nhng phn b phc tp hn, d trn lc a hay trn bin cng u
khng thnh tng bng. Trn bin, tr s ca i kh p cao ln hn mt cht so vi gn
xch o. vng dng nc m v vng dng nc lnh d cng mt v , cng
chnh lch nhau 2 - 3 ln. Mt khc, trn lc a nu lng nc trong t y , th
nhit to ra do bc hi ch yu quyt nh bc x thun. Cn nhng vng sa mc,
na sa mc, t thiu nc, th gn bng vi lng ma nm. Lng bc hi ln nht
ton nm lc a c th n 100 mm ( cao ct nc), trn bin c th n 200 mm.
6
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

7

(c) Lng trao i nhit dng xoy trong nm (kcal/cm
2.
nm)
Hnh 4.2 a, b, c. S phn b a l trong nm v cn bng lng nhit (kcal/cm
2.
nm)
(Buuko, 1956), phn gch xin l thiu ti liu
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Lng trao i nhit do dng xoy (trong hnh 4.2c du m v dng l xc nh
ngc), tt c mt i lc v phn ln mt bin u cung cp nhit cho khng kh, tr s
c nm ca vng sa mc v nhit i l ln nht, t 50 - 60 kcal/cm
2
nm tr ln.
Bin i trong nm v cn bng
lng nhit
a)
1 2 3 4 5 6 7 8 9 10 Thng
R
LE
H
Hnh 5.2 l s bin i trong nm
v cn bng lng nhit ca mt s a
im thuc cc vng kh hu in hnh
(trn hnh v ngoi bc x thun ra, du
ca cc s hng khc xc nh ngc).
Hnh 5.2a biu th tnh hnh ca
thnh ph H Ch Minh vng kh hu
gi ma xch o, bc x thun ca ma
kh cao, ma ma thp (i xch o ni
chung, tr vng gi ma ra, bin i
trong nm ca bc x thun rt nh).
5.2a. Vng gi ma xch o (Tp. H Ch Minh
10
0
47' v Bc, 52
0
59' kinh ng)
b) Nhit to ra do bc hi rt cao vo
u ma kh, thp xung rt nhiu vo
cui ma kh v t kh (thng 4), n
ma ma li tng ln.
LE
H
R
0
1
2
3
4
5
6
K
c
a
l
/
c
m
2

n

m

Ch l tr s bc hi thp nht n
mun hn tr s bc x thun ln nht,
quan h gia lng nc trong t v s
bc hi c th ni r vn ny. S bin
i trong nm v trao i nhit dng xoy,
thng l ngc li vi trao i nhit bc
hi, ma kh rt cao.
1 2 3 4 5 6 7 8 9 10 11 12 Thng
5.2b. Vng kh hu lc a nhit i
(Kraxnvxc, 40
0
v Bc, 52
0
59'
kinh ng)
1 2 3 4 5 6 7 8 9 10 11 12 Thng
-2
R
LE
H
10
4
2
0
c)
Ly vng kh hu lc a l
Kratnvxc Trung lm th d,
nh hnh 5.2(b). y do nguyn
nhn ca thin vn hc, bc x
thun bin i trong nm tng i
ln, m ma ng c tr s m. V
ma t, lng nhit bc hi rt nh,
vo ma h li cng thp. Do ,
trao i nhit dng xoy c bit
cao vo ma h, vt bc x thun
vo ma ng hng t khng kh
xung mt t.
K
c
a
l
/
c
m
2

n

m

5.2c. Vng kh hu gi ma v trung bnh (Vlaivxtc
43
0
07' v Bc, 134
0
54' kinh ng); R: bc x thun,
LE: nhit to ra do bc hi; H: Trao i nhit xoy
Hnh 5.2a, b, c. ng cong bin i trong nm
v cn bng lng nhit cc vng kh hu
in hnh (Buuko, 1956)
8
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Ly Vlaivxtc lm th d v vng kh hu gi ma c v trung bnh (hnh
5.2c, tr s ca cc loi cn bng lng nhit ma h y chu c ch ca tri rm, nn
ng cong hi bng.
trn bin ni chung, trao i nhit dng xoy ht sc nh, bin i trong nm
cng rt nh. Bc x thun v nhit to ra do bc hi khc nhau theo v v vng kh
hu, v lng nhit to ra do bc hi cao hn hn hoc thp hn hn bc x thun m
sinh ra thiu hoc tha lng nhit; tnh trng thiu hoc tha ny c b p bng
lng nhit i lu gia tng su v tng nng ca bin hoc lng nhit vn chuyn
nh dng nc bin.
1.2. Cn bng nc trn ng rung
Da vo nh lut bo ton nng lng, c th dng cng thc cn bng lng
nhit biu th tnh hnh phn phi li nng lng mt tri chiu trn ng rung.
Cng l lun nh vy, c th dng cng thc cn bng nc ni r s phn phi li
nc trn ng rung. Cng thc cn bng nc ca ng rung cho thy l ch trong
mt thi gian nht nh, tng lng nc dng rn, dng lng, dng hi m khng
gian chung quanh cung cp cho ng rung v lng cc loi nc mt i phi bng
khng. Cng thc nh sau (Buduko, 1956):
r + E + fw + m = 0 (10)
Trong cng thc ny r l lng nc ma, E l chnh lch bc hi v ngng t trn
b mt ng rung; fw l nc chy mt trn mt t; m l trao i nc ca mt t
vi tng di ca ng rung. Du ca cc s hng trong cng thc (10) ging nh
trong cng thc cn bng lng nhit, phng hng vo ng rung c tr s dng.
Gi tr ca m bng tng ca nc trng lc t mt t chy xung tng su, nc ca r
thc vt hp thu v lng lu ng theo chiu thng ng ton b nc trong cc tng
c hm lng khc nhau. Cng thc 10 cng c th dng cho trng hp bin i t
nhiu, tc l nc lu ng theo chiu thng ng bng tng lng nc trong t chy
ra fp v hm lng nc trong t tng mt b.
Tng ca nc b mt chy ra fw v nc trong t chy ra fp bng tng lng
nc chy ra f (f = fw + fp), do biu th thnh cng thc sau y:
r + E + f + b = 0 (11)
Cng thc ny c th dng tnh ton cn bng nc ca c mt ci h, hoc cn
bng nc ca mt vng nht nh (th d lu vc mt dng sng). Lc ny, f l tng s
nc phn phi li theo chiu nm ngang trong thi gian nghin cu nht nh, c mt
nc v trong tng y t. Nu ly tr s bnh qun th s hng b rt nh. Cng thc
(10) c th bin dng thnh cng thc sau y:
r + E + f = 0 (12)
i vi ton qu t, phn phi li ca nc theo chiu nm ngang bng 0 vy:
r + E = 0 (13)
Bnh qun nm ca vng t sa mc khng ti s nh cng thc (13).
9
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

2. Mi trng t
t
t c cy trng hoc thc vt bc cao sinh trng pht trin trong h sinh thi cn
gi l t trng hay th nhng. Cng vi cy trng, t cng l i tng kinh
doanh ca con ngi. Nhng cho t mt nh ngha chnh xc tht khng phi n
gin nh vy. Thnh phn ch yu cu thnh t c nhin l nhng cht v c trong
m, nhng nu khng c v s vi sinh vt, ng vt sinh sng trong t v cht hu c
phn gii t xc thc vt, ng vt, th khng th coi l t. Cho n nay, nh ngha
v t c nhiu ngi tha nhn l nh ngha ca acutraiep. Kira (1959) cho rng
t l th kt hp khng th chia ct ca sinh vt v v sinh, do h thng tc dng v
phn tc dng cu thnh, v ch ra rng Ci tr thnh mt b phn ca h sinh thi no
, lu di ti cng mt a im, trong tc dng ln nhau vi gii sinh vt, tr thnh
ci c cu trc nht nh thch ng vi nhng c tnh ca h sinh thi y, tc l
t. t nng nghip chu rt nhiu nh hng ca con ngi, kt qu l lm cho tc
dng ln nhau ca sinh vt v v sinh c thc y hoc b c ch. Chi tit v nhng
pht trin gn y ca mn hc t, tc dng ln nhau gia t v cy trng, ngh
tham kho nhng ti liu lin quan, y ch ch yu ni r vn hnh thnh t lin
h cht ch vi mi trng vt l.
Thnh phn ca t
t c ba pha, u tin l pha rn, tc l phn th rn bao gm nhng mnh vn,
cc thnh phn v c ca sn phm phong ho m v cc cht hu c t l sn
phm phn gii xc sinh vt; ri n pha kh v pha lng nm gia khe hng ca pha
rn. gi l khe ba pha ca t. T l ca pha rn, pha lng v pha kh, cng tc l s
phn b ca ba pha, d l cng mt loi t cng thay i, nht l trong iu kin kh hu
khc nhau, t l pha lng v pha kh thay i kh ln. ng rung, do cy ba v cc
bin php canh tc, s phn b ba pha ca t cng khc nhau. Ni chung, s phn b ba
pha ca t do s khc nhau v chng loi t v v tr lp t m hnh thnh tr s c
tnh tng ng.
Thnh phn ca pha rn, theo to nh ca ht m chia ra ct, limon v st. S hp
thnh theo ng knh ht ca nhng ht cht v c ny gi l thnh phn c gii. Phn
loi da theo gi l phn loi t theo thnh phn c gii. V d: t ct pha, t tht
nh, t tht trung bnh, t tht nng, t st nh, t st trung bnh v t st nng.
Trong thin nhin, cc ht ct, limon v st thng t dng ht n m chng
thng lin kt vi nhau nh cc keo hu c v v c to thnh cc ht c kch
thc ln hn. Ht kt trong t c hnh dng khc nhau tu theo loi t: dng phin,
dng tr, dng hn, dng cu v cc bin th ca chng. Cc ht kt ny c th xem nh
cc vin gch b nh cu trc thnh t. Nhng t c cu to t cc ht kt vin
c gi l t c kt cu vin hoc t c cu trc vin. l loi t c ph
thin nhin cao, nh t en vng n i hay t bazan ca Vit Nam.
10
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Nc trong t
Nc trong t dng lng c th gi l dung dch t, v n cha cc cht ho tan
gm nhiu loi cht v c v cht hu c. Cn c vo lc lin kt ca nc trong t
vi ht t, c th chia: nc lin kt cht vi hp cht khong, nc ht m, nc lm
nho, nc mao qun v nc trng lc. Trong , nc m cy trng c th ht l
nc mao qun v nc trng lc, gi l nc hu hiu. biu th cng ht nc
trong t trn ht t, Schofield (1935) ngh dng logarit biu th cao ct nc
(cm) tng ng vi lc ht gi l PF. Hnh 6.2 ni r quan h ca PF vi loi nc
trong t v hng s nc trong t.
PF
Kh p
tng ng
Loi nc
trong t
Hng s ca nc trong t
Phng php
o
Dch c vn
Nc trng lc
Nc mao
qun
Nc ht m

0



1,5


2,7




4,2

4,5
6,0
7,0

0,001



0,031


0,5




15

31
1.000
10.000 Nc kt hp
c vn
Dung tch lng
Lng gi nc ln nht
Gii hn trn tnh do, im
thnh ht
Lng gi nc ng
rung
Lng gi nc nh nht
ng lng nc im
kh ng mao qun
Gii hn di co rt
Gii hn di tnh do
im kh ho
Kh m mt thy c
H s ht m (di m bo
ho)
H s ht m (R.H - di 50%)
105
0
C kh mt nc

Hnh 6.2. Quan h ca PF, kh p tng ng, loi v hng s nc trong t
v phng php o
- H s ht m: Nc ht t khng kh m vo khi ri mng t ra.
- H s kh ho ( m cy ho): Lng nc trong t lm cho thc vt bt u
ho gi l h s kh ho ban u, lng nc m sau khi ho khng th phc hi li
nguyn trng gi l h s kh ho vnh cu.
- ng lng nc: Cho t bo ho nc, a vo my ly tm tng ng vi
1000 ln trng lc, nc cn li trong t l ng lng nc, gn tng ng vi
nc mao qun.
- Lng gi nc ng rung: Nc ma v nc ti tr thnh nc trng lc di
ng xung di, sau i ln nh tc dng mao qun, khi loi nc ny hu nh ngng di
11
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

ng, lng nc ca tng t mt, gi l lng cha nc ng rung. Tr s PF khong
1,5-1,7. Trong khong gia ca tr s ny v h s kh ho ban u l nc hu hiu.
Hnh 6.2 cn cho bit phm vi c th ca cc phng php khc nhau o nc trong
t, cn c vo phm vi ca tr s PF cn thit m chn phng php o tng ng.
Khng kh trong t
Thnh phn khng kh trong t cng ging nh kh tri, gm xi, nit, cacbonic
v cc kh him khc. im khc nhau ch yu gia khng kh trong t v khng kh
trong kh quyn l hm lng CO
2
. Trong khng kh thng thng, hm lng CO
2

khong 0,33% cn trong khng kh tng t mt thng l 0,2 - 1%. Trong rung nc,
c th khng kh ho tan vo nc mt rung ri khuch tn vo t. Trong t, oxi
c tiu dng, sinh ra CO
2
, H
2
v mtan, thnh bt kh i ln mt nc ri vo khng
kh. Thnh phn ca khng kh trong t s d khng ging vi khng kh thng thng
l v s h hp ca r thc vt v vi sinh vt cn tiu hao oxi v thi ra CO
2
.
Bng 1.2. S tiu hao oxi trong t khi c cy trng v khng c cy trng
Lng tiu hao oxi (l/m
2
.ngy)
t Cy trng
Tng lng
tiu hao
Khng c
cy trng
Lng tiu hao
cho cy trng
t ct pha
t than bn
Khoai ty
Thuc l
7,6
13,0
4,8
9,4
2,8
3,6
Bng 1.2 ni r s khc bit v lng tiu dng kh oxi, trng thi c cy trng v
khng c cy trng. Bng chng r rng l oxi tiu hao cho sinh vt t nhiu hn cho cy
trng. Trong t gn b r, s tn ti ca b r thc y hot ng ca vi sinh vt, cho nn
trong thc t, oxi dng cho cy trng cn nh hn na; cn v ng thi CO
2
trong t s
c cp ti phn sau.
3. Mi trng sinh vt
Sinh vt trong t
Nhiu loi ng vt v thc vt c tr trong t. Trong thc vt ch yu l: nm, vi
khun, x khun, to; ng vt c loi bin hnh amip, b hung, ng vt tit tc ln, giun,
ng vt thn mm... Nhng sinh vt t ny trong qu trnh chuyn ho nng lng ca h
sinh thi ng rung, l loi tiu dng v loi phn gii nng lng, lin h vi nhau khng
qua tc dng v phn tc dng ca h thng ch th - mi trng (hnh 7.2). V vi sinh vt
t ngh tham kho gio trnh Vi sinh vt t ca Trng i hc Nng nghip I.
Cn trng, sinh vt ngun bnh
S hin din ca cn trng trong sn xut cy trng thng c coi l c hi, hon ton
tri ngc vi h thng c nh, chuyn dch nng lng mt tri ca cy trng. V th trng
im nghin cu thng l phng tr su hi. ng v gc ca h sinh thi ng rung, li
rt ch n vn sinh thi ca nhng qun th ng vt, t ra cng phi lm r s chuyn
ho nng lng tun hon vt cht ca nhng qun th ng vt ny v sinh vt ngun bnh.
12
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

13
C di
C di trong h sinh thi ng rung l i tng c nhng nh nghin cu cy trng
v nhng nh sinh thi hc thc vt ht sc quan tm. C di l i th cnh tranh ca cy
trng, l i tng phi phng tr. Gn y trong vic nghin cu c di, ngy cng c nhiu
ngi vn dng phng php sinh thi hc (c l ngi u tin i theo hng ny l Arai,
1961). V c di trong h sinh thi ng rung, s c cp n mc cnh tranh..

Bn phn
Thuc dit c
Thuc dit nm
Thuc dit su
Ma axt
Chn v to ging
Canh tc
Cc loi cng sinh Cc loi giun, ng vt t c tc dng
phn hy cc cht hu c
Loi k sinh nm, virus
Cc loi cng sinh
Loi n c
Loi hoi sinh
Loi n tht Loi k sinh
Hnh 7.2. Mi quan h gia cy trng, cc loi sinh vt t, cn trng di
cc bin php iu khin ca con ngi thng qua h sinh thi ng rung
(Ngun: Shiyomi v Koizumi, 2001)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

14
4. Cu trc ca qun th cy trng
Cu trc sn xut ca qun th cy trng
Bt k cy trng no cng u cu thnh do nhiu phin l quang hp tc l h
thng quang hp v h thng khng quang hp nh r, thn. Hn na, v cy trng trn
ng rung l qun th do nhiu c th hp thnh, cho nn quang hp v sn xut vt
cht ca mt phin l so vi mt c th hoc thm ch mt qun th, th tnh cht v
mc phc tp ca n rt khc nhau.
Khi so snh sn xut vt cht ca mt phin l vi mt c th, ch yu phi xt
trong quang hp, do phng thc xp t phin l khc nhau nn mi phin l nhn
c cng chiu sng s khc nhau, nn cng quang hp cng khc nhau. Hn
na, trong vic phn phi v tiu dng sn phm quang hp, cng phi xt t l s
lng gia h thng quang hp v h thng khng quang hp, t l hnh thnh h thng
khng quang hp... S xp t ca l, hnh thi ca tn cy c s khc nhau gia cc
loi thc vt, iu ht sc quan trng i vi s tm hiu quang hp ca c th v
qun th. Hnh 8.2 cho thy, khi din tch l bng nhiu ln mt t (khi l ht sc rm
rp), l nm ngang khng c li cho tng quang hp ca tng l.

Hnh 8.2. M hnh xp t nhm l. Gi thit din tch
lp trn v di nh nhau, nhng hng b mt v v tr
tng i khc nhau. Nu nh sng chiu t trn xung,
th lng ng ho ca lp l di l ln hn


Ngi ta gi tnh trng xp t l nh vy l h thng ng ho, nhng cha tin hnh
phn tch nh lng h thng ng ho.















Hnh 9.2. Hai loi hnh cu trc sn xut ca qun lc trong h sinh thi ng c
(Monsi, Saeki - 1953)
A. Cy du giun - Chenopodiun album L. qun lc thun.
B. C voi - Pennisetum purpurascens Nak. qun lc thun, phn mu en l loi khc.
Hai loi hnh cu trc sn xut ny cng thy qun th cy trng.
Lng sn xut
ca h thng
khng ng ho
Lng sn xut
ca h thng
khng ng ho
Lng sn xut ca
h thng ng ho
Lng sn xut ca
h thng ng ho
B- Loi hnh l hp h Ho tho
Cng chiu sng
A- Loi hnh l rng
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Ngi u tin ng v gc sn xut vt cht, nu r c nh lng cho cu trc
qun th thc vt l Monsi v Saeki (1953). H tin hnh chia tng ct cy theo nhng
khong cch nht nh t cao xung thp ton b thc vt trong mt din tch nht nh, ri
phn bit nh lng theo h thng ng ho (h thng quang hp) v h thng khng ng
ho, tm ra s phn b theo chiu thng ng s lng cc tng, gi l phng php ct tng.
S phn b v lng ca cc tng tm c nh hnh 9.2, bn tri ng gia l h thng
quang hp, bn phi l h thng khng quang hp, lm thnh bn cu trc, sn xut. Trong
hnh v ly cng chiu sng b mt qun th l 100, bn trong qun th ln lt biu th
cng chiu sng theo cao, s khc bit theo tng l ca loi hnh l rng v loi hnh l
hp h ho tho, iu kin chiu sng v cu trc qun th c quan h r rng, c th thy
ngay c. Hnh v ny tuy l hai loi hnh in hnh cu trc sn xut ca qun th ng c,
nhng v c bn cng c th phn nh loi hnh cu trc sn xut ca qun th cy trng.
Phng thc biu hin cu trc qun lc ly phng php ct tng v bn cu trc sn
xut lm c s ch nu r c sinh khi ca tng l v h thng quang hp, nhng cha xt
n s xp t tng l nh Boysen - Jensen nu ra, y l mt vn cn phi nghin cu.
H thng l v h thng thn ca qun th cy trng
Sumia (1960) dng phng php nghin cu sn xut vt cht, tin hnh phn
tch ging nng sut cao ca cy trng. ng pht trin thm mt bc cch suy ngh
v h thng ng ho ca Boysen - Jensen, gi tng l ca c th hay qun th l h
thng l. Phng php tin hnh phn tch h thng l ca Sumia l (1) xem chiu
hng ca l, tng phin l tri bng hay ng thng; (2) t l din tch trng lng ca
phin l (t din l), tc l dy hay l mng; (3) din tch ca mt phin l to hay nh;
(4) trng thi xp t (nm ngang) ca l, chia ra loi hnh tha v dy. Trong khi phn
tch h thng l, ng thi quan st mu l m nht, hm lng m... ng v gc
sinh thi ca ging, phng php ny khng dng li giai on nh tnh, m
tin mt bc vo nhng nh lng, v mt nghin cu sn xut vt cht cng cn kh
nhiu ni dung nhng cha nu c r rng quan h gia n vi cu trc mi trng.
Cu trc sinh hc ca qun th cy trng
Ross (1970) xut pht t lp trng xng nghin cu vt l hc i vi qun
th thc vt, tin hnh nghin cu t m cu trc qun th (cu trc tng l).
Ross v Ninson (1965) cho rng: phn tch nh lng qun th cy trng, ch
cn tm ra hm s phn b mt din tch b mt ca cc c quan thn, l, bng v
hm s xp t khng gian ca chng. Din tch b mt ca l cng tc l din tch l,
thng ch l mt na ca tng din tch, cng l din tch mt mt ca l. Ngoi ra, i
vi thn v bng, khi nghin cu lt nh sng (mc thu quang) bn trong qun th,
c th ly din tch mt ct. Khi tm hm s phn b mt din tch b mt cc c
quan khc nhau, cn gi thit l cc c quan xp t tu c trn phng nm ngang. Gi
thit ny thch hp vi qun th pht trin bnh thng, cn thi k u pht trin cha
th c c. Do , hm s phn b mt din tch b mt ch xt n s bin i theo
chiu thng ng l c. C ngha l, i vi l ch cn tm ra din tch l c n v
th tch khng gian trong phm vi cao z. Vy c cng thc sau y:
15
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

16
dz (z)
r
U
h
0 0
L =
(14)
dz (z)
L
U
h
0 0
L =
L
0
: ch r din tch l
H: cao ca qun th (cm)
U
L
(z): mt din tch l (cm
-1
) cao z (cm).
Cng nh th, c th tm ra ch s din tch b mt ca thn nh sau:
(15)
Cn v hm s xp t ca l g
L
(z) th phn tch nh sau: trc ht coi phin l l
mt tm phng, l cong hoc nhn c th coi nh mt phin l c chia nh ra thnh
nhiu tm phng v coi cc tm phng nh mt phin l. Sau , ly phng ch mt
trn (b mt) ca phin l l phng ca php tuyn dng trn mt l, nh vy th php
tuyn ca l tri nm ngang s l thng ng, php tuyn ca phin l ng thng l
nm ngang, phng chiu ca php tuyn ) r (
L
do hai to sau y quyt nh:
)
L , L
0 ( )
L
r (

= (16)

L
: l gc thin nh ca php tuyn trn mt l (gc hp thnh do php tuyn v
trc thng ng o c t pha trn dng);

L
: l gc phng v ca php tuyn (gc hp thnh do php tuyn v trc Bc -
Nam, ly Bc l 0
0
, tin hnh o thun theo chiu kim ng h).
Trong qun th cy trng, trn thc t c th coi b mt l hng v na trn qu t,
khng gian ca php tuyn trn mt l l 2;
L
trong khong
2
0

;
L
bin thin trong
khong 0 ~ 2. S phn b ca php tuyn trn mt l cao z tu , biu th nh cng
thc sau u:


=
2
) , 0 , z ( g
2
) r , z ( g
L L L L L
(17)
Vi mi phng ca
) r (
L
tin hnh tch phn i vi g
L
, s thnh:
1 d sin ) , , z ( g d
2
1
d ) r , z ( g
2
1
L L L L L
2 /
0 L
2
0 L L L 2
=

(18)
Trong :
d
L
= sin
L
d
L
d
L
, biu th n v gc lp th chung quanh
L
r , phng tu .
Hm s g
L
) r , z (
L
biu th t l chim ca din tch l trn php tuyn trong ca n v
gc lp th phng chiu
L
r so vi ton b din tch l trong n v khng gian cao
z; v gi l hm s sp xp khng gian ca l. Gi thit
L
khng c quan h vi
L
, th:
g
L
(z;
L
,
L
) = g
L
(z;
L
) g
L
(z,
L
) (19)
Vn thng dng hm s sau y thay cho g
L
(z;
L
) :
g
L
(z;
L
) = g
L
(z;
L
) sin
L
(20)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

17

Trong trng hp ny:
(21) 1 d ) , z ( g
L L L
2 /
o
=

(22) =


2 d ) , z ( ' ' g
L L L
2
0
Trong : g
L
d
L
l t l chim ca din tch l gc thin nh ca php tuyn
gia
L
v
L
+ d
L
so vi tng din tch l.
t gi thit gia
L
v
L
khng c quan h tng quan, ni ng ra l khng
c y , nhng ch tm gi tr gn ng th c th b qua quan h tng quan. Ross
ng v gc ca hnh hc khng gian tng l biu hin cu trc qun th cy
trng nh vy v gi n l cu trc hnh hc ca tn cy.
Warren Wilson (1965) ngi xtrylia, dng phng php im mu phn tch
cu trc tng l. ng gi gc ca l, din tch l chia tng l stand structure (cu trc
ca thm cy). W.A. Williams v ctv. (1968) M cng dng phng php ging nh
vy tin hnh nghin cu tng l v gi n l canopy (community architecture - cu trc
tng tn ca qun th). V sau, c mt s ngi cng dng cc danh t ny, nhng cha
c s ng h ca nhiu ngi.
Phng php o cu trc hnh hc qun th cy trng
Phng php o bng thc o nghing l: Theo phng php ct tng ca
Monsi v Saeki, khi tin hnh ct theo cao nht nh, dng thc o nghing l
ca Laisk (1965) o gc thin nh ca php tuyn ca mi phin l (thc t l bng
gc to thnh bi mt l v mt nm ngang),
L
chia v ct theo rng nht nh tu ,
ln lt tm din tch l. Ly din tch l tm c chia cho tng din tch l ca tng ,
s thng l g
L
(z). Th d, t
L
chia v ct theo gc 15
0
th nh s th t 0
0
15
0
,
15
0
30
0
... l j = 1, 2 ... v c ng thc sau y:

= = =
J
1 j
LJ j
LJ j
LJ j L
) , z ( ' g
) , z ( g
) , z ( ' g ) z ( g (23)
y
)
, (
LJ J
z bi
c tuyn c
' g u th tng din tch
a l m php gc thin nh l
ch
uan h vi gc
ph
Hnh 10.2. Thc o nghing l
LJ
c cao z. Th c o nghing l do
Laisk thit k nh hnh 10.2.
Dng bn phng v quyt nh hm
s sp xp l g (z, ) c q
L L
ng v ca php tuyn trn mt l: do gc
phng v ca php tuyn, ln lt tm din
tch theo nhm php tuyn ging nh trng
hp gc thin nh, tnh ton bng cng thc
sau y:
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

18

k L L
g
) , z ( ' ' g
) , z ( g

k
1 k
Lk k
Lk
k L k
) , z ( g
) , z (
2
(24)
y, g
k
'(z,
Lk
) biu th tng din tch ca l m php tuyn c gc phng v
Lk

cao z
in tch l tch ring theo gc thin nh v gc phng v ca nhng php tuyn
khc
.
D
nhau c tin hnh o ng thi, nn i vi mt s tng cn o s nhm j k din
tch l. Lc ny c quan h nh sau:

= =


=

k
1 k
k L Lj jk
j
1 j
Lk Lj jk
k L k j L j
) , , z ( g
) , , z ( g
) , z ( g ). , z ( ' ' g
2
(25)
g
Jk
(z,
Lj

Lk
) biu th tng din tch ca l m php tuyn c gc thin nh
LJ
,
gc ph
(cn gi l phng php qu tch)
o t
bt u t phn chn
l, c

c
sinh trng pht trin. Tnh hnh bin i
gc nghing ca nhm l biu hin rt r.

L , L L
) , z ( g
ng v
Lk
cao z.
Phng php hnh chiu
Hnh 11.2 cho thy, trn ng rung
Hnh 11.2. Phng php hnh chiu
Hnh
Z
3
Z
2
Z
P
1
P
2
P
3
X
X
Z
nghin cu qun th cy trng
ny ly cy ng lm v d. Trn t th
nghim, o to X - Z ca P
1
, P
2
v P
3
ca
mi phin l, dng chiu di phin l v
qu tch ca l
o X - Z ca phn chn l (P
1
),
im cao nht ca phn cong (P
2
), nh
(phn nhn) l (P
3
) ca mi phin l ca
cc cy. o ton chiu di l ca l, v qu
tch ca l ln hnh v. Sau , ct rng
z tu , dng thc o gc o gc
nghing
L
ca l trong tng z v z + z,
lc ny chia ct l ra mt cch thch ng,
cho n mc c th nhn thnh tm phng
(trn thc t, mt phin l ng c th chia
ct theo 5 - 10 cm).
o din tch l
10cm o mt ln, dng phng
php hnh hc o din tch ca l.
Cch lm sau ging nh trng hp
dng thc o nghing ni trn.
Hnh 12.2, bn v hnh chiu tm
bng phng php hnh chiu phn
nh s bin i ca cy ng trong qu trnh
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

19
Phng php im mu
cm vo qun th theo gc nht nh, ghi chp s
ln c, chia cho khoang cch m kim cm vo, s tm
thy


300







200
100





0




Dng chic kim nh di,


g tng l m mi kim cm
c tn tip xc. nghing bnh qun ca tng l tnh bng cng thc sau:

00 00
:
f
0
l s lng cm c khi kim di ng nm ngang 10cm;
0
l s lng cm c khi kim di ng thng ng 1 cm.
Sau khi tm
F = f
00
sec (27)
Ngy 14 thng 7 Ngy 25 thng 7 Ngy 9 thng 8 Ngy 20 thng 9 (1955)
Hnh 12.2. Bn v hnh chiu tng l ng (ging lai s 7)
bin i qua cc thi k sinh trng pht trin
0 0
f
f
157 , 0
f
f 1 , 0
2
tg =


= (26)
Trong
f
0
c th mt din tch l F l:

Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

20
ng nh t
din tch l theo tng. Hnh 13.2 l th d o mch en nhiu hoa (Lolium
mult
2 l kt qu o cu trc hnh hc thi k chn ca qun th i mch. C
th t n; tng trn cng v tng
thp
0








0
1
2
3
4
5
6
0
1
2
3
4
5
6
1 2 3 4
(
c
m
)


c
a
o

0.0 0.2 0.0 0.2 0.0 0.2 0.4 0.6 0.0 0.2 0.4 0.6 0.8
F F F F
G ing l (F) (cm
2
/cm
3
)
S tun sau khi gieo
c ngh ca l ( ) (), mt din tch


0
1
2
3
4
5
6
0
1
2
3
4
5
6
0.065 0.209 0.966 1.919
Ch s din tch l
ng
ng php
0.007 0.050 0.447 0.852
Ch s din tch l chiu thng
Hnh 13.2. Th d v o cu trc tng l bng ph
im mu



















o theo t vy c th tm c gc nghing bnh qun ca l v m
thc khi
iforum).
Trng thi thc t ca cu trc qun th cy trng
Hnh 14.
hy l: din tch b mt ca bng i mch tng i l
nht, l c nghing nh chim t l ln; l dng ng (70 ) tng gia kh nhiu.
Ngoi ra, gc phng v ca l ta nh phn b tu c. Hn na, i vi i mch v
hng dng, gia mt trng v cu trc qun th c quan h nh hnh 15.2.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

21
Bng 2.2. Phn b l theo gc phng v
Nam Ty Bc ng
La m
Ng 0,23 0,27 0,24
0,25
0,26
0,26 0,26 0,23

thi k u sinh tr
ca cch x l m a cc cch x
l kh c nhau cng p xp ca l c
quan
oss (1970). a s l quay theo mt tri, sng sm
hn
m c theo gc nghing mt l lm hm s phn b
ca
4 loi hnh nh hnh 17.2.
0
0,4
0,2
60 - 80 cm

g
J
(
w
)

0






100
80




ng, p n tch hn b mt di l r rng l chu nh hng
t , n thi k sau, c bit l i mch, nh hng c
khng r rt, hu nh khng c khc bit. Hm s s
h vi gc nghing, gc phng v ca l, v thi k sinh trng pht trin c bin
i rt ln, nhng khng thy nh hng ca cch x l mt i vi n. iu c
ngha l hm s sp xp ca l l c tnh c hu ca loi hoc ging, kh c th ci
thin qua phng php trng trt.
Cu trc tng l ca mt s cy trng thay i trong ngy r rng. Nhiu cy h
u nh u tng, l c vn ng trong ngy, l ca hng dng cng nh vy. Hnh
16.2 l kt qu nghin cu ca R
g ng, bui tra hng nam, bui chiu hng ty. Nhng vn ng ca l chm
30
0
so vi vn ng ca mt tri.
Wit (1965) dng phng php ging nh ca Ross, tm ra hm s phn b
ca l. V gc phng v, ng dng kt qu tu c ca Nichiporovic (bng 2.2) v ly
s phn b tng i din tch l t
l. Wit chnh l kt qu o c cc thi k sinh trng khc nhau ca
chiu cy trng nh ng, ci ng, c ba l trng, khoai ty, mch en... v chia ra lm
60
40
F=3.20


c
a
o

(
c
m
)

40 -60 cm
0,4
0,2
0
20 - 40 cm
0,4
0.2
20
0
1 0,04 0,0 ,12 0,16
M din tch b mt l
(cm
2
/cm
2
t)
8 0
t

0
10 - 20 cm
0.4
0.2
0
0 15 30 45 60 75 90
g ca l Gc nghin
0 90 180 220 360
Gc phng v ca mt l
0
0,2
0,1
0
0,2
0,1
0
0,2
0,1
0
0,2
0,1
0
g
K
(
w
)

4 u trc hnh hc ca qu i mch
Bng
l
Thn
Gc
n th .2. C Hnh 1
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

tn l
) l
ng,
i ty.
Tn



22












Trong hnh 17.2,
ngang (pla-nophile canopy
cu trc ca loi hnh l b
nh c ba l trng v khoa
l ng (erectophile
canopy) l cu trc ca loi hnh
l ng thng, nh mch en.
Tn l nghing (plagiophile
canopy) l ca loi hnh l gc
nghing 30 - 60
0
chim a s,
nh ci ng, ng, ci du, kh
nhiu cy trng thuc loi hnh
ny. Nhng Wit cha ch ra
loi tng ng vi tn l bng -
ng (extremophile canopy).
Ngoi ra, cy c mch en lu
nm (Lolium perenne) ma xun
l loi hnh l ng, sau khi qua
mt v ct, t loi hnh l ng
thng tr thnh loi hnh l
nghing, n ma h li tr
thnh loi hnh l bng.

G
30 60
c ng ng ca l
L
m

s


p
h

n

b


90
hi
H

0,5
1,0
1,5
3
(
4
1
2
7
8
b)
0
4
5
6
a)
L
)
0,1
0,3
6 8 10 12 14 16 18
Gi
0,2
H

m

s


p
h

b


1
2
3
4
4
2
1
c)
Hnh 16.2. Bin i trong ngy c tr hnh hc
nhm cy ng d (Ross, 1970)
a) Bin i trong ngy ca gc nghing mt l:
1: 1 - 6 gi 30 pht; 0 g


chia 10
0
;
3
S
W
N
E
0,5
a cu c
l h ng
2: 1 i 30 pht
3: 12 gi 30 pht; 4: 16 gi 30 pht
b) ng cong t l ca din tch l (gc nghing mt l
theo 10
0
mt) trong tng din tch l: 1: 0
0

2: 10
0
- 20
0
........... 7 : 60
0
- 70
0
; 8: 70
0
- 90
0
.
i mch
Dy
Va
Tha
0
20
40
60
0 0,2 0 0 0,1 0,1
H dng
7/7 29/7 8/3
0,2
ng
17/7
0
0
30
10
1
2
0,05 0 0,05 0 0,05 0 0,05
Mt din tch l (cm
-1
)
i mch Hng dng
29/1 Dy
Va
Tha
7/7
0
60
00
17/7 20/3
25/4
11/5
8/8
29/7
1 2 3 4 5 6 1 2 3 4 5 6
0
0,2
Gc thin nh php tuyn trn mt l
LI
Hnh 15.2. nh hng ca mt trng u trc hnh hc ca nhm l
0,4
0
0
0,4
0,2
0
,4
0,2
0
0
0,2
,4
n c
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

23
n th cy trng lm ch th ca h sinh thi ng rung, th
ch th - mi trng. nu
r q
h s
trn
ng php
cn
l
ng rung nghin
Bu
rong phng
php
vic nghin cu trng thi kh hu tng khng kh gn mt t.
5. Cu trc mi trng ca h sinh thi ng rung
Nh trn ni, ly qu
mi trng vt l ca n s c xem nh l mt h thng
uy lut vn ng ca h thng ny, iu quan trng nht l lm sng t quy lut bin
i nng lng v vt cht ca n. i vi mi trng vt l th phn tch vt l tng
khng kh gn mt t, cn i vi sinh vt (cy trng) th nghin cu kh hu sinh hc.
Vt l ca tng khng kh gn mt t
Xt v mt kh tng hc, v tr ca
inh thi ng rung trong phm vi
mt t 50 - 60 cm, tc l trong
tng khng kh gn mt t. Do , s
bin i nng lng v vt cht gia cy
trng v mi trng b cc nh lut vt
l c tc dng i vi tng khng kh gn
mt t quyt nh. Mi trng bn ngoi
(iu kin kh hu) c th chia ra dng
nhit v dng nc ca tng khng kh
gn mt t, k c tng t canh tc.
Chng phn nh qu trnh v kt qu trao
i nhit v nc. Thng qua vic nghin
cu s cn bng lng nhit v cn bng
nc trong tng khng kh gn mt t c
th lm sng t cc nh lut vt l quyt
nh cc qu trnh trao i ny.
Nghin cu vt l i vi tng
khng kh gn mt t, ly ph
bng lng nhit v c hc khng
kh pht trin nhanh chng lm
phng php chnh.
Cn bng lng nhit, nh trn ni
cn bng lng nhit
Tn l bng
Tn l ng
Tn l nghing
Tn l bng - ng
0 30 60 90
0
Gc nghing ca mt l (
L
)
n

t

c
h

l


0,8
Hnh 17.2. Lo
loi theo gc nghing mt l (De Wit,1965)
g
Planophile canopy - tn l bng
Erectophile canopy - tn l ng bng thng
Plagiophile canopy - tn l nghin
Extremophile canopy - tn l bng ng
i hnh phn b din tch l phn
0,2
0,4
0,6
T

n
g

i

t

n
h

t
o

n

c

a

d
i

mt phng php da vo phng trnh


cu s trao i hin nhit v tim nhit.
uko nghin cu c h thng i vi vn ny, khng nhng dng cho vic
nghin cu vt l tng khng kh gn mt t, m cn dng cho vic nghin cu nhiu
mt khc nh trng thi lng nhit trong nh knh, nh m trng cy.
Phng php ng lc hc khng kh tc l dng phng php pht trin t c
hc hng khng vo vic nghin cu tng khng kh gn mt t. T
dng xc nh bc hi - thot hi nc, kt hp vi s pht trin l lun v
dng xoy khng kh, tr thnh phng php cn thit khng th thiu c trong
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

24
ng
nng
l k
g lng hin nhit ca dng
xoy n tro t
l n
ng
t
gy ca cng thc (2a)
ng kh xung
mt
Xt n qu trnh mt nhit do bc hi nc ca t, IE phi gim nh dn v H
tng ln, t s ca thng lng hin nhit v thng lng tim nhit l:
Hin nay, mt s ngi lm cng tc kh tng nng nghip khi nghin cu tiu
kh hu ng rung s dng cc phng php .
S bin i trong ngy v cn bng lng nhit ng ru
Hnh 18.2 l mt th d, nu r
s bin i trong ngy v cn bng
Rn = 4.58mm
S = 0.65mm
E = 5.66mm
H = -0.64mm
Rn
E
H
S
00 0400 0600 1200 1600 2000 2400
Gi /9/1957)
1,0
1,2
lng theo cng thc (2).
t qu quan st trn ng c gieo
hn hp Luzec v Bromut c ti
y . Bui tra, phn ln bc x
thun dng cho bc hi - thot hi
nc, thng lng hin nhit nh
trao i dng xoy l khong 15 %
ca bc x thun; ban m v chiu
ti, bc thot hi nc vt bc x
thun, ban m hu nh gn bng 0,
hoc t nhiu cng l s dng.
Trong hnh v, tr bc x thun ra
du ca cc s hng trong cng thc
(2) u ngc li, tc l:
R = IE + H + B (2a)
Lng nhit dng cho thot hi
nc vt xa thun, ngha l
thn
0,8
0,6
0,4
0,2
0
-0,2
00
(4
v nhit truyn d ng
gun nng lng b sung cho
nng lng thiu. Cng nh hnh
18.2 cho thy, kiu phn b nhit
ban m xut hin vo bui ti, nhit
di t trong mt thi gian tng
na bc x thun. Hnh ny cn biu
tc l tr s ca cc s hng ca (2b).
R = IE + H + B (2b)
T bit c lng thot hi vt bc x thun. Mt khc, nhit to t di
t ln mt t, ng thi cng c nhit vi s lng gn nh th t kh
L

n
g

n
h
i

t

l

u

Hnh 18.2. Bin i trong ngy v cn bng
l
n v trn trc tung l
t
2
G
R
ot hi nc)
ng nhit ng c chn nui (Tanner, 1960)
mm/gi.
mm/gi = 0,1 cal/cm . pht
ieo hn hp Medicago + Bromut c ti y
n - Bc x thun; S - Nhit truyn dn trong t
E - Thng lng tim t (bc hi v th
H - Thng lng hin nhit
truyn dn trong t ch truyn t mt t xu
i ngn bui tra, ban m ch bng khong m
th lng tnh ton trong n
t b ch thiu ca bc x thun.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

25
H
=
IE
(28)
T s ny gi l t s Bowen, l
mt
ca t s Bowen sau khi m au
ma, IE gim, H tng ln, t s n
tng
t khng kh, hi nc,
nng
b mt
d
c x ng dy c
ch s din tch l l hiu sng.
100
0
1 2 3 4 5 6 7
S ngy sa
nh 19.2. Bin i t s a
(Rauner, 1960, Uch 1964)
i lng nu r s phn phi
nhit nng trong tng khng kh gn
mt t. Hnh 19.2 nu r s b i in
a. S
Bowe
dn.
Phn b iu kin kh hu trong
v ngoi qun th cy trng
Hnh 20.2 biu th s phn b
theo phng thng ng bc x
thun, nhi
CO
2
v tc gi trong v
ngoi qun th ng v phn
in tch l ca qun th ng.
l kt qu o c t hai qun th c
mt trng khc nhau.














Bc x thun v sut phn x
thun bn trong qun th gim rt nhanh, r rng nht l khu tr
n. iu ny c quan h cht ch vi s gim c
B

10
19 - 22/7/1954
14 - 17/8/1954
11-16/9/1954


B
o
w
e
n

(
L
0

/

I
E
0
)

T


H
0
u khi ma
Bowen sau khi m
ijima v hnh,
60
120
240
180
300
360
420
480
540
600
u

c
a
o

c
m

LAD
LAI 3.6
LAI 2.6
Tc
gi
CO
0 1 2 3 0 80 160 240 5 0 5 8 9 10 20 22 24 26 28 0 2 4 6 8 10
dm
2
/ dm
3
u cm / s c ppm e gm /m
3

0
C Ly / min f,
C
h
i

2

n
nh 2 . Phn b iu kin kh hu trong v ngoi qun th ng mt trng
khc nhau (Lemon, 1970) tr s bnh qun trong khong thi gian
11 gi 45 pht - 12 gi 15 pht ngy 15-8-1968
Ni quan st v
Trc
H
2
O
nhit
Bc x
thu
H 0.2
o: Ellis Hollow, Ithaca, NewYork.
khi o ta cy, iu chnh mt .
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

26
nh 21.2 cho thy s bin i trong
ngy v cn bng phn x rung nc (a(Q +
a), du trong cng thc (3) ngc li). C
th t ng
66%
ch l.
v
thnh ng gch gch trong hnh v.
Mt khc, bc x thun, nh cng
thc (3) cho thy, l hiu ca thnh phn
hn
-
a tun mt, thu thp v tnh
t qu o chiu sng trc x v chiu
, biu th nh hnh 24.2. C th
t
chi
sn

H
hy bc x thun c th t n kho
bc x mt tri. Bc x hu hiu sng
di thng thng l tr s m (t mt t
hng ln khng kh), bui tra tr s gn
nh c nh, bng 0,2 ly/min
(1)
.
Hnh 22.2 cho thy s bin i trong
ngy ca sut phn x ca mt s cy trng,
bt k loi cy trng no, s bin i trong
ngy u r rng, hn na, sut phn x cn
bin i theo ch s din tch l, khi LAI
( ) 2

(ch s din tch l) = 2,5 tr ln th xp x
bng 0,2 (hnh 23.2). Quan h ny biu th bng cng thc sau:
a = a
p
- (a
p
- a
w
) exp (-0,56F) (29)
a
p
l sut phn x khi cy trng rm rp
a l sut phn x ca mt nc,
F l tr s din t
Tr s tnh theo cng thc (29)
g xung chiu sng trc x, chiu
sng tn x v thnh phn hng ln
sng di. N
ton k
sng tn x
hy l, thi k ma phn hu nh khng c
u sng trc x. Ngoi ra, t l chiu
g trc x cao nht c th t 60% tr ln.

( ) 1
Ly/min l n v biu th cng bc x mt tri, tc l s calo chiu theo mt cm
2
mt pht.
( ) 2
LAI = Leaf Area Index; exp = exponent: s m.
0,3
1,0
0
0,5
(Q+q)
R
S
a(Q+q)
0 4 8 12 16 20 24
10/8/1965 Gi
Hnh 21.2. Bin i trong ngy
v cn bng bc x rung nc
8
10
12
14
16
1
2
2
24
26
8
0
2
28
30
32
Khoai lang 12/8
C bc n 1/6
La m 21/5/1969
La nc 5/7
Tr s o
Tr s tnh
ton
0 1 2 3 4 5 6
Ch s din tch l
h 23.2. Quan h gia ch s ch
l v sut phn x run

t

p
h
5 6 7 8 9 10 17 18 19h
nh 22.2. Bin i trong ngy v phn x
ca mt s cy trng
11 12 13 14 15 16

n

x


(
%
)

H

Hn
S
u
din t
g nc
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

27
i:
c gi c s k hng thng trn
ng rung, tc gi bn tro . V phn b tc gi
ca ng, c nh lut logarit sau y:













G
T hc nhau do mt ca qun th cy trng. T
ng qun th gim xung rt nhanh
ng ru
V
*
z - d
u (z) =
K
ln
z
o
(30)
Trong : V
*
= (T/)
1/2
tc ma st (cm/s); K = 0,4 hng s Kaman;
z: t ; d hiu chnh mt t (cm); z
o
: m g
gh : l a st ca khng kh (g/cm
3
).
hng c nh
cao cch m
(cm); T
t
c m
= tr s c
ng
Ni mt cch cht ch hn, cng thc (30) ch c xc lp khi k
hng ca s phn b nhit. Morin v Obuenop ch ra: cng thc phn b tc
gi khi c phn b nhit trong tng khng kh gn mt t, nh sau:
V
*
z - d z - d
u (z) =
K
ln
z
o
+
L
(31)
: 6,0 hng s, L l di Obuenop.
Hnh 25.2 ch h b o a tr s hiu chnh t d
v g gh z
o
tro t c trong rung i mch. Tr hiu
chn gi, khi tc gi t n 4 m/s
tr l
r tnh
ng cng
nh in i the tc
o
gi c mt
s hc (30) v (31)
h mt t gim xung rt nhanh cng vi tc
n th tr thnh mt tr s nht nh, khng thay i trong sut c thi gian sinh
trng. g gh z
o
thi k u sinh trng c tng ln tng ng theo tc gi,
nhng li gim xung vo thi k cui sinh trng. C th v thi k cui sinh trng
do tc gi tng ln, cy bin thnh nghing, d gim i, z
o
tng ln. Mt s kt qu
quan st khc cng ph hp vi kt lun ny.
0
1,0
2,0
3,0
Tng bc x sng ngn (Q+q)
(bc x mt tri)
Chiu sng tn x (q)
Chiu sng trc x (Q)
Chiu sng tn x (q)
Thng Thng Thng
Hnh 24.2. Chiu sn a n
n mt ln
c

x
l
y
p

t
h

n
h

c

n

x
0

t

t
r

i

/
t
h

i

k

t
r

c

x


,4

0
0,2
0,6
0,8
T

c
h
i

u

s

n
g
t
g tn x v t l c
theo tnh ton na tu
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

28
(32)
V phn b tc gi bn trong qun th cy trng, nh hnh 25.2 cho thy, gim
rt nhanh tng trn qun th, ngi ta dng cc cng thc vi hm s m sau y
biu th s phn b tc gi ny:
Cng thc Saito: ) (z p - ( exp u (z) u
H
=
Z
H


dz)

Z

V
*
H
: u (z) =
K
ln
o

= dz )
H
- (1 - exp u (z) u
H
Cng thc Inoue:
(33)
- d
Cng thc Takeda
z


) z H (
F 1
F
2
C
exp
D

n ng, u l tc gi b mt qun
th, H l cao ca qun th cy trng, F l mt cy trng, C
D
l h s khng.
u ni r: tc gi bn trong q
mt qun th, tu theo
h s
xun
CO t
tng mt din li
tng v
70 - 80 ppm; khi F
t

= 4,2 th nng l 100 ppm; c th thy, lng h thp nng CO
2
vo bui tra l tng
(34)
y, p (z), l h s c xt s phn b tr cy
H
Ba cng thc ni trn un th cy trng khc vi b
c trng phn
b mt cy, gim
0
0,4
0 400 800
g thnh hm s
m. Ngoi ra, cng
thc Saito v Inoue
c lp ra theo kinh
nghim sau khi nghin
cu l lun i vi
cng thc Takea,
nhng gn b mt
qun th, th cc cng
thc ny u gn
ging nhau.
Nng CO
2
:
xem hnh 26.2, ta thy
b mt qun th nng
nhanh, gn
2
tng dn ln, cn bn trong qun th th gim xung r
tch l ln nht th thp nht, cn n gn mt t th
ln. y ni r,
pha di, n gn mt
Hnh 26.2 cho thy s bin i trong ngy ca nng CO
o bui tra, CO
2
ca ng ung lu ng t pha trn x
t th ngc li, t mt t lu ng ln trn.
r ung
2
b mt qun th cy
ng. C th thy r, nng CO
2
gim vo bui tra; trong trng hp ch s din tch l
0,5 tr xung, nng l 20 - 30 ppm. Nu ch s din tch l (F
t
tng ln, tc l khi F
t
=
0,56 th nng CO
2
tng ng l 30 - 40 ppm, khi F
t
= 2,5 nng l
0,6
Tc gi (U
150
) (cm/s) Tc gi (U
150
) (cm/s)
0 400 800
0
0,04
0,08
0
(b) Z
0
/h
Hnh 25.2. Tr s hiu chnh t v g gh trong
cng thc phn b tc gi bin i theo tc gi
(a): qu t (d) v
(b): qu cao
U 150 t t 150 c ch
0,08
(a)
0,04
mt
an h ca tr s hiu chnh mt
an h v
cao cy (h) vi tc gi
cy (h) vi tc gi. ca t s g gh (z
0
)
l tc gi ch cch m m. Kt qu rung i m
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

29
ln tng ng. S bin i ny r rng l c quan h cht ch vi quang hp. S bin i
nng CO
2
ca ngy 8 v ngy 9 thng 8 tuy ni l thi k sinh trng pht trin ging
nhau, cng vn c khc nhau r rt, nguyn nhn chnh c th l tc gi trn qun th
khng ging nhau, tc gi ch cao cch mt t 245 cm bui tra ngy 8 thng 8 l 40
cm/s, cn ngy 9 l 50 - 90 cm/s.
S phn b theo phng thng ng ca nng CO
2
c th tnh theo cng thc:
P
H
z - d
C
o
- C
z
=
kV*
ln
z
o
(35)
C
o
, C
z
l nng CO
2
(g C
3
O
2
/ cm ) cao d + z
o
v z; p
H
l thng lng CO
2

trn
Nu p
H
(tng ng vi cng
p ca qun th) l nht
nh, th nng CO
2
t l nghch vi V*.
bn trong qun th, nh hnh 26.2 cho
thy
thn
xung thp n cao 100 cm (cch mt
chn

25 ng
nh hi cao hn khu trng cy. iu ny c
hnh 27.2, b mt qun th, nng ny gim dn
theo
a r, s ht nc v cht dinh dng. nhit t ch yu do lng nhit m
mt t tip thu (lng nhit truyn dn xung di t) v tnh cht nhit ca t quyt nh.

thm thc vt (g CO
2
/ cm
2
.s).
25, 26/7 F
t
=
135cm
8, 9/8
245

quang h

tr s
Di y s cp ti nng CO
2

, cng c bin i trong ngy r rng.
Nht l ngy 8 thng 8 tc gi tng
i nh, quang hp mnh, c th hin
h mt ct rt pht trin. Tng thp ca
nng CO
2
, sng sm xut hin gn
tng mt qun th, sau v tr
( ) 3
ca n
thp xung theo vi mt tri ln cao, n
11 gi 35 pht, khi gn gi ng Nam
( ) 4
,
t). Lc ny CO
2
trong v ngoi qun th
nhit khng kh v nhit t nh hnh
hnh 26.2 c th thy nhit nhit hnh
th c quan h vi tc gi bn trong
qun th.
Quan st nng hi nc
h lch n 27 ppm.
nhit, m: dng thc phn s
.2. T s phn b nhit khng kh tro
cao. C ngha l ngc li vi s phn b CO
2
ni trn. Vo bui tra, hi
nc lu ng t mt t ln trn, tc l s bc hi v thot hi nc rt mnh.
nhit t ca ng rung c nh hng rt ln n s ny mm ca ht ging, s
pht trin c

( ) 3
Tc cao cch mt t
( ) 4
Gi ng Nam l gi mt tri i qua ng Bc - Nam
2,5
205cm
26
25
15, 19/6
F
t
=4,2
8
9
cm
F
t
=4,2
20/8
F
t
=0,0
18
19
25cm
30/6 F
t
=0,05
30cm
70cm
15
14
14, 15/7
300
250
25
300
350
0
250
300
150
300
150
300
300
Hnh 26.2. Bi i trong ngy ca
CO
2
mt qun th ng
Hnh ny biu th ch s l F
t
ao
s cong l ngy thn ng trn o. Ch
v c
g o
din tch
nng
n
b

Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

30
khu
khc gi l c trng
dn
i, s lu thng nhit
di
Cc i lng c trng cho
nhit ca t l: h s truyn dn
nhit ca t (cal/cm.s.
0
C) h s
2
ch tn nhit ca t K (cm /s),
dung lng nhit ca t C
V
= c
(cal/cm
3
.
0
C).
Trong bn i lng trn, v
C
V
l cc i lng c trng c bn,
cc i lng
xut.
Hnh 28.2 biu th s bin i
trong ngy, bin i trong nm ca
nhit t v s lu ng ca nhit
tng ng (nhit truyn dn di t).
Nh trn n
t c th tnh bng cng thc
(8), trn thc t
0 z
dz
s
dT
=
kh
xc nh, nn thng dng ba phng
php
cng thc sau:
sau y:
Phng php phn tch nhit:
cn c vo s phn b theo phng
thng ng ca nhit t, tnh bng
n T . C . n . v
n
(36) H dt B
t
n
r
=


H
t
1 n
0
1
=
ca tng t
n
: dy
khc nhau (cm);
n . v C : dung lng nhit
bnh
t
1
)n
Cng thc (36) cho thy,
ly c
mi tng t
phn,
o
qun ca tng t khc
nhau.

n
= (t
2
-
hnh lch nhit thi
gian t
2
v t
1
ca
nhn vi dy tng
t v C
n
bnh qun ca cc
tng, tng ca cc tng tc l
B .


c
a
o

(
m
)

0
6
0
5

0
7
0
5

0
8
0
5

0
9
3
5

1
0
3
5

1
1
3
5

1
2
3
5

1
4
0
5

1
5
0
5

1
6
0
5

1
7
0
5

1
8
0
5

1
9
0
5

8/9/1966
2
3
8/9/1966
0
7
0
5

0
8
0
5

0
9
3
5

1
0
3
5

1
1
3
5

1
2
3
5

1
4
0
5

1
5
0
5

1
6
0
5

1
7
0
5

1
8
0
5

1
9
0
5

0,02
0,02
mg/l


c
a
0
0
3
4
o

(
m
)

2
1
1
Nng CO
2
0,04 0,02 0
Mt din tch l (cm
-1
)
Hnh 27.2. Bin i trong ngy ca nng CO
2

trong qun th ng (Uchijima v ctv, 1968)
Ch s trn ng cong l thi gi n gch bn phi hnh
biu th phn b phng thng t din tch l

an, ph
ng m s
30
Bin i
trong ngy
1 cm
300cm
Bin i
trong nm
10 cm
20cm
100cm
20 cm
50 cm
900cm
Tr
nhit To nhit Tr
nhit To nhit
-40
0
40
0
20
N
h
i

t

(
0
C
)
y
0
-0,2
0,1
10
N
h
t

(
0
C
)
i

L
y
/
n
g
a
y
0 2 16
Gi
1 6 8
Thng
2 2 4 10 12 20 24 4 8 1
H
c
nh 28.2. Bin n i trong n
a nhit t ng nhit di t tn g
m
ng
i trong ngy, bi
lu v
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

(T - T )
Phng php tm truyn nhit: Vn dng phng php o lng nhit truyn qua
tng, cc nh kin trc dng cng thc sau tnh ton trong k thut truyn nhit:
a e u
B
z
=
L
= a (T
e
- T
u
) (37)
T
e
v T
u
l nhit (
0
C) mt trn v mt di ca tm truyn nhit

a
l h s t nh cal
L l dy ca tm (cm); a l hng s thc nghim
e
- T
u
.
huyt dn nhit m tin hnh x l
n ong nm) ca nhit mt t,
tm
ruyn it ca tm ( /cm.s.
0
C)
Trn thc t, bn trong tm truyn nhit c kp cp nhit in, dng o T
Phng php truyn dn nhit: Da vo l t
g cong biu din s bin i trong ngy (bin i tr
0 z
s
dT
= ca cng thc 8, v:
dz
dTs c

dz
z = O = T sin (t +
4
+ ) (38)
0
D
31
o : )
2
t sin( T . ) t , O ( B
o
+ c

+ = (39)
T bin bin i nhit t.
gc thi gian, gc lch pha.
V quan h gia bin bin i nhit mt t v s lu ng nhit di t,
biu
s k
hnh thnh
hn

0
l
= 2 /
th nh hnh 29.2.
Phng php xc nh h
huch tn dng xoy v
tc trao i:
Quan st khi t ng khi
i ra, c th thy s lu ng
ca khi khng
g c quy tc. Li nh khi
gi bo, ng h o tc gi
v ng h o hng gi cng
khng ngng dao ng. Cho
nn s vn ng ca khng
kh khng phi l s lu ng
c quy tc, m l khng quy
tc, cng tc l dng xoy.
Gi l dng xoy, nh trong
cng mt th long, bao gm
mt cch quy tc nhiu ch
0 4 8 12
Bin bin i nhit t (T
0
)
0
C
16
mt
B
O
Q
S
120
80
40
0
160
120
80
40
g

n
g

O
n

nh 2 . Quan h ca bi bin i t
m t
160

t

(
Q
t

l

u

S
)

T

n
g

n
h
i

t

l

n
g

l

n
g

t
r
o
n
c
a
l
/
c
m
B
i


b
i

i

t
r
o
n
g

n
g

y

n
h
i
t
r
o
n
g

t

(

B
)

l
y
/
m
i
nhi

2
.

n

a

n
g

y

H 9.2 n
t t v lu ng ng nhit di
(Uchijima, 1961)
Q
S
l tng nhit lng tch tr trong t trong sut
thi gian nhit t ln cao
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

32
lc: Khuch tn dng xoy cao z
1
v z
2
trn b mt
qun
(40)
(lng khuch tn dng xoy), C l gam
CO
2
trao i (cm/s) gia z
1
v z
2
;
z
1
v z
2
.
xoy phc tp to nh khc nhau. Nhng ch xoy nh vy gi l dng xoy. Trong tng
khng kh gn mt t, do s c st ca khng kh vi mt t hnh thnh nhng lu
ng c th, gi l tng ranh gii dng xoy. Ngoi ra, thng lng vt cht (khuch
tn) trong dng xoy so vi thng lng tng khng dng xoy th ln hn r rng.
Bi v thng lng (lng khuch tn) c dc nng vt cht v h s khuch
tn dng xoy quyt nh.
Phng php kh ng
th cy trng, ly CO
2
lm th d, c th biu th nh sau:
P = D
1 - 2
(C
1
- C
2
)
Trong cng thc ny, P l thng lng
/cm . s;
D
2
1-2
l tc
C
1
, C
2
l nng CO
2
(gam CO
2
/cm
3
) ch
D
1-2
l s nghch o ca h s khng khuch tn r
1-2
Quan h gia chng nh sau:

= =

2
1
z
z
2 1
2 1
) z ( K
dz
1
r
1
D
(41)
K l h s khuch tn dng xoy (cm
2
/s). T cng thc (40) v (41) c th thy,
nu

bit mt s no trong ba s: h s khuch tn dng xoy, h s khng
khuch tn v tc trao i, th c th da vo lng chnh lch nng vt cht
trong tng khng kh gn mt t (CO
2
chng hn) tm ra lng khuch tn dng
xoy. ng dng cng thc phn b tc gi (30) ni trn, th ng lc ma st ca
khng kh - lng khuch tn dng xoy ca lng vt cht vn ng tnh bng cng
thc sau:

2
1
2
2
1 2
2
d z
d z
ln
) u u ( K


=
(42)
Cn c vo cng thc (41) v (42), tnh tc trao i ca lng vt cht vn ng
nh sau:

2
1
2
2
1 2
2
d z
d z
ln
) u u ( K

=
(43)
Thay cng thc (43) vo cng thc (40) c th tm ra thng lng CO
2
(lc ny gi
thit trong s lu chuyn lng vt cht vn ng v lu chuyn CO
2
c h s khuch
tn dng xoy bng nhau). Phng php da vo vic xc nh phn b tc gi
tm h s khuch tn dng xoy, ri mi tm thng lng CO
2
nh vy, l phng php
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

33
, dng cng thc
(43)

do Thornthwaite v Holzman xut trc tin (1941) tm thng lng hi nc,
sau mi c gi l phng php ca ng lc hc khng kh.
Hnh 30.2 cho thy: da vo tr s Uchijima o c trn ng c
tm c tc trao i, bin i trong ngy ca h s khuch tn dng xoy, tc
gi tng ng (U
200
) v tc ma st. Bui sng gi to, tc ma st, tc trao
i v h s khuch tn dng xoy u tng ln, trong tc trao i theo st nht
vi s bin i ca tc gi. Trong tit ni v s phn b tc gi nu ln, gi l
nh lut logarit, ni mt cch cht ch th ch khi tng khng kh gn mt t l trung
lp (tc l khi khng c s phn b nhit) mi c xc lp. Do , cng thc (42)
v (43) rt ra t cng thc (30) rt ra t cng thc (30) s khng dng c khi tc
gi nh, trong tng khng kh c s phn b nhit r rt (nh ma ma h...). Xt n
c s phn b nhit th tc trao i c th biu th nh sau:


















2
1
2
1 2
2
) 2 1 (
d z
d z
ln
) u u ( K
) i R . 1 ( D


(44)
y i R gi l s Richardson bnh qun, biu th nh cng thc sau:
0 0 1
0 5
0
1
0
0
1
5
0
3 4 5 6 7 8 0 1 2
0-200
/s)

K
2
0
0
D
U
2
0
0
Tc gi U
200
(cm/s)
Tc trao i D (cm
2
0
0
4
0
0
6
0
0
8
0
0 2 3 4 5
4
8
1
2
2
0
6
2
4
4
V
*
Tc ma s V
H x
0
0
3
N
g

Hnh 30.2. Bin i trong ngy ca h s khuch tn dng xoy, tc trao i,


tc ma st v tc gi, (U200 l tc gi ch cao 200cm)
y

o
:

1
3

-

1
4
/
8
/
1
9
5
3

(x1 tn oy
cm
2
/s)
K
20
khuch s
*(cm/s) t

1
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

34
2
2 1
i
T g
R

=
u
T
T
l hng s kinh nghim, g l
gia t
, (44)
cn
d v
ca nhit v hi nc (K
H
, K
W
) trong tng
khn v z
R -B
D
1 - 2
=
I (
(45)
Khu trng dy
420
Khu trng tha
cao cy trng
360
c trng trng (cm/s
2
), T
1
v T
2

l nhit khng kh cao z
1
v z
2

v tr s bnh qun ca chng (dng
nhit tuyt i
0
K); z = z
1
+ 0,5 (z
2

- z
1
); u biu th tc gi ca z.
Khi s dng cng thc (43)
mt vn l xc nh d. Nh
ni trn, d bin i theo tc gi,
nn trc ht dng cng thc phn b
tc gi khi trung lp (tc l khng
c phn b nhit) lm r quan h ca
Phng php cn bng lng nhit
300
240
180
120
60
0


c
a
o

(
c
m
)

5 1x10
4
5 1x10
5
5 1x10
6
5 1x10
7
5
2
/mi
Hnh 3 xoy
tr
K - h s khuch tn (cm n)
1.2. h s khuch tn dng
ong v ngoi qun th cy ng (cm
2
/min)
tc gi.
Gi thit h s khuch tn dng xoy
g kh gn mt t l bng nhau, th tc trao i gia cao z
1 2
biu th nh
sau:
1+ )
Trong cng thc ny,
a
= nh lch m tuyt i (g H
2
O/cm
3
) ca z
1

v z
it rt mnh,
ln
Rz - B
d 1 dq
a
1
- a
2
l ch
2
. Cng thc (45) ly cng thc cn bng lng nhit (2a) lm c s, dng t s
Bowen, H = . IE, lp ra. Do , phng php ny c gi l phng php cn
bng lng nhit. Khi tm tc trao i bng phng php ny, do mt tri lc hin
lc n bc x thun gn nh bng khng, sai s c th ln, nn phi ch .
Bn cnh qun th, hot ng hp thu v to ra hin nhit v tim nh
g bc x thun cng bin i nhanh, cho nn cng khng th dng cng thc (45).
Uchijima da vo cng thc cn bng lng nhit biu th h s khuch tn dng
xoy cao no trong qun lc (K
z
) nh sau:
Kz =
- Cp (
C
)
(46)
dz
+
p dz
Hnh 31.2 biu th u ch tn dng xoy trong v ngoi qun th ng do
Lemon (1970) tm c theo s liu ca hnh 26.2. Khu trng tha trong qun th v
khu trng dy trong tng khng kh gn mt t biu hin tr s tng i ln.
h s kh
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

968).
th c
khu vc rung thong. Ni n gi nguyn trng thi, pha trn hng
gi c
it - m k kiu thng gi, b
phn
Trong cc phng php nh gi h s khuch tn dng xoy, ngoi phng php
c hc khng kh v phng php cn bng lng nhit nu, cn c phng php
cn bng lng vn ng v phng php hm s cu to ca dng xoy (Inoue, 1
Cch quan trc ca phng php kh ng lc hc v phng php cn bng
lng nhit
Khi tin hnh o bng phng php trn cn ch : tng kh trong v ngoi qun
y trng phi gi c nguyn trng thi ca ng rung c quan trc, tin hnh
quan trc trn
n t nht c khong cch 50 - 100 ln o cao o, ni c iu kin tng i
tt cn c din tch khong 1 hecta. Hnh 32.2, l mt v d dng c o lng khi xc
nh thng lng CO
2
bng phng php cn bng lng nhit.
Bc x thun: o bc x thun trn qun th dng bc x k thun Beckmann, o
bc x thun bn trong qun lc dng 4 chic bc x k thun Fank, treo trn dy thp,
di ng qua li trong khong 7 m ly s bnh qun.
Ly mu CO
2
: c cc l ly mu trn thp ly mu khng kh cao 4m, dng bm
ht ly.
nhit, m: l ly mu khng kh c lp nh
cm ng l cp nhit in, dng minivon k ghi.















35


Hnh 32.2. B tr thit b xc nh h s khuch tn bng phng php cn bng nhit
1.B c x thun Becman; 2. My o chiu sng; 3. My o phn x; 4. Thp ly mu kh; 5. L ly
m kh; 6. Dy thp di ng bc x k; 7. Bc x k thun Frank; 8. My o tc gi dy
nhi t; 9. My phn tch kh; 10. Milivon k biu bng in t; 11. Tm truyn nhit; 12. My o

chiu sng kiu ng; 13. Bm chn khng kiu cnh qut; 14. Milivon k kiu bm nt; 15. Milivon
k kiu t ghi; 16. Bn v mt ct ca thp ly mu khng kh gn l ly mu khng kh 5 c lp b
phn cm ng nhit ca nhit k kiu cp nhit in.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Nhit truyn dn trong t: su khong 1 cm c lp tm truyn nhit, dng
minivon k ghi.
aeki (1953) xut m hnh ton hc quang
hp in cu sinh thi sn xut vt cht ca
qun
nhn
Chiu sng v phn x: Chiu sng trn qun th dng my o chiu sng
Consinke, bn trong qun lc dng my chiu sng kiu ng, v sut phn x dng my
o phn x.
Khi quan trc bng phng php kh ng hc, pha trn qun lc cn t nht 5 - 6
ci ng h o tc gi, c kh nng th tt nht nn dng 7 - 8 ci, lp ln lt theo
bi s cao t di ln trn, 4 ci thnh mt nhm, chun b 2 nhm.
6. Quang hp ca qun th cy trng
M hnh quang hp ca qun th cy trng
M hnh nh sng: T khi Monsi v S
ca qun th thc vt cho n nay, vic ngh
th thc vt ly quang hp ca qun th lm trung tm, c nhng pht trin rt
ln. Cn c ca nhng nghin cu ny l s chiu lt ca nh sng trong qun th v
ng cong quang hp ca phin l, do c th gi l m hnh nh sng v quang hp
ca qun th. V m hnh nh sng s trnh by thm phn sau.
M hnh hiu sut nhn nh sng: Mt s nh nghin cu cy trng nh Murata,
Takeda ly vic nghin cu quang hp ca la nc lm trung tm, nghin cu
quang hp v sn xut vt cht ca nhiu loi cy trng. Nghin cu ca h ch yu
bng phng php phng th nghim. V quang hp qun th P, c th biu th nh sau:
P = Afp
0
= Ap
0
e
-bA
(47)
Trong : p
0
l cng quang hp ca mt phin l: A - din tch l; f - hiu sut
nh sng, b - hng s thc nghim.
Mt s nh khoa hc xut khi nim hiu sut nhn nh sng do tch ca
quang hp l n v din tch l quyt nh. Nh cng thc (47) cho thy, hiu sut nh
sng gim vi hm s m ngc theo s tng ln ca din tch l, tuy tng i gn st
vi mc lt nh sng ca qun th, nhng do khi nim c phn cha r, nn cha
lm cn c cho l lun. Ta nh l mt m hnh ca nh sng, k thc vn l m hnh
hiu sut nhn nh sng.
M hnh khuch tn: c mt s nh kh tng hc nng nghip nghin cu vt l
tng khng kh gn mt t ng dng phng php c hc khng kh vo vic
nghin cu s quang hp ca qun th. Cng c ngi thng qua vic nghin cu thng
lng (khuch tn) CO
2
trong tng khng kh gn mt t, m y mnh vic nghin
cu s quang hp qun th.
Nhng suy ngh c bn v m hnh ny s ni sau y: Gi thit trn mt cnh
ng rng, cy trng tin hnh quang hp mnh. Quang hp cng mnh th tiu dng
cng nhiu CO
2
gn mt l cy trng, nng CO
2
s cng h thp. b p vo s
h thp , s cn vn chuyn CO
2
t trong khng kh bn trn qun th v t trong t
v mt l. Lc ny do din tch ng rung rng, nn khng phi xt n s lu ng
theo phng nm ngang ca CO
2
. Bi v phn bt k ca dng kh CO
2
, n v thi
36
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

37
s chng khuch tn t l nghch vi nhau:
gian, n v din tch mt ct ca thng lng ca n cng u bng nhau. Do , s
lu ng nh vy ca CO
2
c th hiu l cung cp cho s quang hp ca qun th s
dng. Cch nhn nhn nh vy l khc vi m hnh ca nh sng, c th gi l m hnh
khuch tn.
M hnh chng khuch tn: M hnh chng khuch tn c th coi l bin th ca
m hnh khuch tn, ch yu do cc hc gi chu u xng, cho rng quang hp
qun th P v
[CO
2
] air - [CO
2
] cl
P =
[r
a
+ r
s
+ r
mc
] CO
2
(48)
[CO
2
] air v [CO
2
] ch g CO
2
trong khng kh v trong
dip lc t, r
a
, r
s
, r
mc
t n ng khuch tn i vi CO
2
ca k gia
khng kh xung quanh v mt l, kh kh t l. Quan h gia s chng khuch
tn v
nhit, h s khuch tn, ... trong v ngoi
qun
; K l h s
thiu
t l
ng
ng tng ng l nn
g l s ch
ng, m th
hong
h s khuch tn nh cng thc (41).
M hnh quang hp qun th cy trng: S quang hp ca qun th c quyt
nh bi hm s quang hp, h hp ca l n (nh sng, nhit, nc), cu trc qun
th v tnh trng phn b CO , nh sng,
2
th. Do , mt s m hnh nu ra y l khng ton din.
S phn b nh sng trong qun th cy trng:
Monsi v Saeki (1953) dng cng thc sau y biu th s lt nh sng vo
trong qun th thc vt ( y l qun th cy trng).
-HF
I = I
o
e (49)
Trong : I
0
l chiu sng nm ngang ca mt trn qun th; I l chiu sng
nm ngang c ng mt qun th a tng F din tch l tnh ton t tiu tn nh sng.
Cn ch : S gim yu ca nh sng l tr s o ca nh sng tn x. Sau ,
Kuroiwa v mt s ngi khc chia nh sng mt tri lm nh sng trc x v nh sng
tn x, ra mt cng thc c xt n cu trc hnh hc ca qun th. y ch gii
nghin cu ca Ross (1970).
V vn gim yu nh sng trc x, Ross v Nilson (1965) ra mt cng
thc c xt n cu trc ca qun th:

L o L
2
L L 0 s 0
d L r r cos ) r , w ( g
1
) w ( U
1
sin ) w ( U
w
cos
r r r
(50)

2
Trong cng thc:
0
- gc thin nh ca mt tri; - t l din tch ca phn
hng v mt tri; w- su t tng mt qun th tr xung; U
g
, U
L
- mt ch
b mt ca thn, l; g
L
(w,
din t
L r ) l hm s sp xp ca l - cng thc (17).
cos
L o
r r

= cos
0
sin
L
+ sin
0
sin
L
+ cos (
0
-
L
) (51)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Trong : ) , ( r ), , ( r
L L L 0 0 0
biu th hng ca bc x mt tri v php
tuyn trn mt l. Da vo cng thc (50) v (51) c th tm ra t l din tch l phn
hng v mt tri tng t k trong qun th (t l din tch m nh s b ng trc x c
th c
L
r
) o thc qun th u tng, tr s tn hiu vo). Hnh (71) biu th g
L
(w, h
c bng cng thc (50) v t l lt vo ca nh sng trc x o c bng cc o
din v mt tri. V nh s tch hng ng tn x, c cng thc kinh nghim nh sau
(Ross v Bichele, 1968):
D (L, h
0
,
L
r
) = D
0
(h
0
) [1 - V (L) sin
2

L
]e
- kdL
(52)
Trong cng thc, D
0
(h
0
) l bc x tn khng gian pha trn qun th, kdL, V(L) l
hng s kinh nghim.
Quang hp qun th biu th bng m hnh nh sng
Quang hp thun A ca phin l n bng tng quang hp p tr i h hp r, gia
quan nghi c trnh
biu
bI
A = - r (53)
g hp qun th v cng nh sng I, thng qua thc m phng
din quan h nh sau:
1 + aI
Trong a v b l hng s thc nghim.


38















0
0,2
0,4
0,6
0,8
1,0
13 thng 8
Kitamoto
14 thng 8
Seigen
0,8
1,0
0
0,2
0,6
0,4
1,0
0
0,2
0,4
0,6
0,8
1,0
26 thng 8
Kitamoto
15 thng 8
Seigen
S thc (
nh 33.2 l lt vo ca nh sng trc x c trong qun th u tng
c theo cu trc hnh hc (Ito v Uagawa, 1971)
S

n
h

t
o


(
0,2
0,4
0,6
0,8

)

0
)
o H . T
v tnh
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

39
y I trn mt l su F tm c hng s cng thc (49) thay vo cng thc
(53), th:
b (KI
0
e - KF)
A =
1 + a (KI
0
e - KF)
- r (54)
L
Quang hp thun ca ton b cc tng ca qun th tm c bng cch ly tch
phn t F = 0 n F i vi cng thc (55):
b
0
P =
Ka
0
)- rF
1 + aKI
ln(
1 + a KI e - KF
(55)
chnh l m hnh ton hc m Monsi v Saeki ni ti. Ross v Bichele cng
vi phng php ging nh vy tm ra tng quang hp K
0
ca n v din tch
phn hng v mt t a din tc ri c h l su L:
1
g (
L
r ) d

K =

0
2
2
1 1
m
+
a [S | cos
L
r r
0
| + D (L, h
0
,
L
)]
(56)
Trong cng thc: a v m l thng s ca ng cong nh sng - quang hp; D (L,
h
0
,
L
) l c .
Mt khc na, quang hp n v din tch l phn rm l:

ng nh sng tn x
K
s ca
1
g (
L
r ) d

K
s =
2

2
1 1

+
(57)
m a D (L, h
0
,
L
)
su L biu hin bng din tch l tnh ton ca b mt qun th, t l din tch
l phn hng v mt tri y l :

s

s
(L,
0
r
) = [ - exp LG (
0
r ) c ec os
Trong cng thc, G bi h c a din tch l hng v
hn
h
0
] (58)
u t l ng h nh hiu hu hiu c
g
0
r . Do , su L, lng quang hp tnh ton l
K
trong tng n v dy
c th biu th nh sau:
0

K
(L, h
0
0
) =
K
(L, h
0
) d L (60)
c (60) v th
quan
) = s K + (1 - s)
K
s (59)
Tnh tch phn i vi cng thc (59) l tm c lng quang hp ca ton th
qun th.
(L
0
, h
Hnh 34.2 cho thy kt qu tnh ton bng cng th i gi it cng
g hp ca l n l nht nh: (1) qun th cu thnh do l thng ng; (2) qun th
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

40
cu hnh o l n ngan v (4 m
l n
i gn 1,5, sau gim xung rt nhanh; t 1,0 ly/min
tr x
h tr quan h
ca
n
s p
y tin hnh trn din tch ng khong 1
ha v
n th tnh t m
ele, 1968)
bc x mt tri, LAI = 6
( 0
0
1. Qun th l thng ng; 2. Q n th l nm ngang; 4. Qun th l
nm ngan
th nh do l phn b u; (3) qun th cu t d m g ) qun th
m ngang chim u th.
T trong hnh (a) c th thy, ng cong ca qun th l nm ngang tng t vi
l n, qun th l phn b t 0,2 ly/min tr ln th gn thnh ng thng. Qun th l
thng ng t tr s ln nht kh
ung th qun th l nm ngang t quang hp cao nht. T hnh (b) cho thy: quan
h ca tng quang hp vi din tch l, trong qun th l tng i tha (ch s din tch
l < 2), quang hp qun th tng rt nhanh theo s tng ln ca ch s din tch l, nh
hng ca cu trc qun th nh hn v quang hp ca loi hnh l nm ngang cao. Khi
ch s din tch l = 2, quang hp ca qun th khng c quan h cu trc hnh hc ca
qun th. qun th c ch s din tch l ln, nh hng ca cu trc hnh hc ln,
quang hp ca loi hnh l thng ng cao nht.














N n cho thy, c im ca m hnh nh sng l ch c th nu r
quang hp qun th vi cc iu kin chiu sng, vi cu trc qun th, c bit l
cu trc hnh hc. Ngoi ra, cn thit phi ch ra rng: dng m hnh ny cha xt
hn b phng nm ngang ca nh sng bn trong qun th (cch gii quyt xt
n s phn b hng nm ngang ca nh sng, ngh tham kho sch sinh thi hc
rng trong ton tp nghin cu sinh thi hc).
Dng m hnh khuch tn nghin cu quang hp ca qun th: Inoue v ctv.
(1968) dng phng php cn bng lng nhit tin hnh phn tch quang hp qun
th, i vi qun th ng. Nhng nghin cu n
theo cc thi k sinh trng pht trin khc nhau ca cy ng. Vic b tr cc
dng c quang trc nh hnh 32.2 ni r.
Hnh 34.2. Quang hp ca qu heo hnh nh sng
(Ross v Bich
(a) Quan h ca quang hp qun th v
b) Quan h ca quang hp qun th v ch s din tch l, cao mt tri 5
un th l phn b u; 3. Qu
g chim u th; 5. ng cong nh sng - Quang hp ca l n
0 0,5 1,0 1,5 Q
0
0 15 30 45 60 75 h
o
Bc x mt tri (Q
0
), cao mt tri h
o
a

q
u

n

t
h


2
90
C

n
g


q
u
a
n
g

h

p

c

m
g

C
O

/
d
m
2

/
g
i


0 1 2 3 4 5 6
Ch s din tch l
a

q
u
2
i

n

t
h

n
g


q
u
a
n
g

h

p

c

m
g

C
O

/
d
m
2

/
g
1
3
4
5
3
4
2
1
2
100
40
80
80
120
a) b)
60
40
20
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

ung, tc gi rt thp. Nhng ngy 20,
tc
Bin i trong ngy ca quang hp qun th: Da vo kt qu bit, chng ta
hy xem lng CO
2
v tc trao i (hnh 35.2). Ngy 8 thng 8 t bui sng n
khong 1 gi chiu, tc gi 30 cm/s tr x
gi bui sng l 200 cm/s tr ln, sau gim nh dn, nhng vn cao hn 100
cm/s. Chiu sng ca hai ngy u rt cao, tr s cao nht chng l 1,2 ly/min. Bin
i trong ngy v tc trao i trong ngy v tc gi c xu th gn ging nhau.


7 9 11 13 15 17 19
Gi (ngy 8 thng 8)
5 7 9 11 13 15 17 19
Gi (ngy 20 thng 8)
0,5
1,0
I
H
U
245
D
225-255
P
h
Pa

i
0
20
30
10
I
H
L
y
/
m
i
n

0

U
200
D
250-300
h
Pa
41
























5
100
0
0
2
4
0
0
-1
-2
0
0
0
1
2
0
0
0
2
m
/
s
C
m
/
s
P
C
O
2
/
c
m
.
S
2
%
1
0
3
g

T
Hnh 35.2. K t qu quang hp o c ca qun th ng bng phng php cn bng
- Chiu sng: U
2
245cm
Hnh 36.2. CO
2
thi t t ra
(Inoue v ctv., 1968)
1 - P
0
= 0.11 x 10
-8
x 3.0
T/10
u
tn

lng nhit (Inoue v ctv., 1968)


I
H 45
- tc gi cao
P
h
- Dng kh (thng lng) CO
2
t tng khng kh gn mt t i ln,
Pa - Dng kh CO
2
t mt t ln,
i
- hiu sut li dng nng lng chiu sng, E
T
- h s bc hi
8
1966
26/7
8/8
9/8
2 - P
0
= 0.15 x 10
-8
x 3.5
T/10

ng cong trong hnh l kt q
h ton bng 2 cng thc trn
19/8
20/8
1
2
C
O
2
/
c
m
2
.
S

0
0 25 30 35
nhit b mt t (
0
C)
6
4

t

t
h
o

t

r
a
2
2
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

42
goi ra, bin i trong ngy v thng lng CO
2
t tng khng kh gn mt t
ln qun th (P
h
), tr on sau bui chiu ra, cng rt ging vi s bin i trong ngy
ca t
onteich (1964)
tm
- nhit t, Q - hng
s th c
ng, c th tm c bng cng thc sau y:
nh
biu
phn b
tn



N
c trao i. Nh trc ni, trong qun th cy trng v c CO
2
to ra t t
ra, nn nng CO
2
gn mt t tng i cao (hnh 36.2). Ging vi trng hp ca
tng khng kh gn mt t, cn c vo tc trao i trong qun th v chnh lch
nng CO
2
c th tm c thng lng CO
2
t mt t i ln qun th cy trng.
Thng lng CO
2
(P
S
) tm c biu th nh hnh 36.2. Trong t sut ngy u thi
ra CO
2
, nhng bui tra cao hn so vi sng, P
S
trc v sau bui tra bng khong
20% ca P
h
CO
2
t t thi ra (h hp t) c quan h cht ch vi s h hp ca vi sinh
vt t v r, y da vo nhng ti liu quan trc c, biu th mi quan h gia
lng thi CO
2
v nhit t lin quan cht ch vi n nh hnh 36.2.
nhit t trong hnh l tr s nhit b mt t. Khi nhit mt t t 25
0
C
ln 34
0
C, P
s
ln khong 4 ln. ng cong trong hnh l quan h m M
0
c trong phm vi nhit 3 - 15 C, o thc t tm ra:
P
S
= P
SO
Q
T/10
(61)
Trong cng thc: P
SO
l thng lng CO
2
khi T
S
= 0
0
C , T
nghim.
Dng phng php cn bng lng nhit nh gi P
S
c th ni l tng i
tho ng.
Quan h ca quang hp qun th vi chiu sng v tc gi: Quang hp thun
ca qun th
P
N
= P
H
- P
S
(62)
Quan h ca P
N
tm c
vy chiu sng c
th nh hnh 37.2.
Mt s im trong hnh kh
phn tn, nhng vn
g i u hai bn ng
cong. ng cong v theo tnh
ton ca cng thc Monsi v
Saeki ni r kt qu ca cng
thc m hnh nh sng v
phng php cn bng nhit l
nht tr vi nhau v i th. Tr
s quan trc v trn hnh s d
F = 4,2
p

t
h
u

1
n


40
C

n
g


q
u
a
n
g

h

1
0
3

C
O
2
/
c
m
2
.
S
0 0,2 0,4 0,6 0,8 1,0 1,2 1,6
Bc x mt tri ly/min
1,4
30
20
10
0
1966
8/8
9/8

19/8
20/8
Hnh
th ng tm
37.2. Quan h gia quang hp thun ca qun
c bn bng lng g phng php cn
nhit vi bc x mt tri (Inoue v ctv., 1968)
ng cong trong hnh l v tr s tnh ton theo cng thc m
hnh nh sng. 1,0 x 10
-8
g CO
2
/cm
2
.S = 3,6mg CO
2
/dm
2
.h
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

kh phn tn, mt l do nhng sai khc v ngy thng o gy nn, hai l cng c nh
hng ca s bin i tc gi. Hnh 38.2 (Yabuki v Hamotani, 1971) biu th thng
lng CO
2
o bng phng php kh ng hc, c th thy r rng l cng quang
hp thun tng theo tc gi tng. l mt im c li ca phng php kh ng
hc.
Cn bng CO
2
ca qun th ng nh hnh 35.2 v tr s ca cc ngy o khc trong
cng thi gian c chnh l li nh bng 3.2.
Bng 3.2. Cn bng CO
2
ca qun th ng ban ngy (6 gi n 18 gi)
(Inoue v ctv., 1968)
P
h
P
s
(P
h
- P
s
) P
s Ngy
thng
Lng chiu sng
(ly/ngy)
mg/cm
2
. ngy (P
h
- P
s
)

I

T
26/7 407,5 - 4,13 + 0,85 - 4,98 0,171 0,031 51,2
27/7 455,1 - 4,88 + 0,92 - 5,80 0,159 0,033 70,0
8/8 528,5 - 5,73 + 1,56 - 7,28 0,214 0,035 92,6
9/8 208,8 - 5,45 + 0,64 - 6,09 0,105 0,074 42,1
19/8 276,5 - 6,06 + 0,82 - 6,88 0,119 0,063 49,7
20/8 450,7 - 7,07 + 1,83 - 8,90 0,205 0,050 47,4
P
h
P
s
tng ng l thng lng CO
2
ca ng rung canh tc v t t thi ra;

I
- hiu sut li dng nng lng ;
T
- h s thot hi nc.
Bng ny ch gii hn ban ngy, trong thi gian ny lng CO
2
t khng kh
chuyn xung qun th l 4,13 - 7.07 mg CO
2
/cm
2
, lng CO
2
t mt t chuyn ln
qun th trong khong 0,64 -1,83 mg CO
2
/cm
2
. Nh vy, s lng t mt t cung
cp chim 10 - 20% tng lng CO
2
m qun th ht. Tr s ny c bn ging vi tr s
do Monteith (1964) tm c.
Hiu sut s dng nng lng v h s thot hi nc: Hiu sut s dng nng
lng bc x mt tri
I
trong quang
hp thun ca qun th tm c nh
vy ghi trong bng 3.2 v hnh 35.2.
Ngy 8/8 tc gi nh, nng
CO
2
thp,
I
l 2 - 4%; ngy 20 tc
gi mnh hn
I
l 4 - 5 %. Nhng tr
s o vo cc ngy khc cng c tr
s cao n 7%, nh vy nu so vi
hiu sut li dng nng lng tnh
theo lng tng khi lng cht kh
th hi cao hn mt cht, c th l do
cha bao gm lng h hp ban m.
43
Mg CO
2
/dm
2
hr
10
20
30
40
12/8/1970
0 100 200
Tc gi U
140
(cm/s)
Hnh 38.2. Quan h ca cng quang hp
thun do bng phng php kh ng hc
vi tc gi (Yabuki, Hamotani, 1971).
Lng chiu sng 1,2 ly/min
0
C

n
g


q
u
a
n
g

h

p

t
h
u

n


1
0
3

C
O
2

/
c
m
2
.
S

Hnh 35.2 v bng 3.2 cn cho
thy r h s thot hi nc (lng
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

44
nc cn) tnh ra bng cng thc sau y:
E
T

T
=
(P
h
- P
s
)
(63)
Trong cng thc:
T
l h s thot hi nc, E
T
l lng bc - thot hi nc tm
c bng phng php cn bng lng nhit, = C
6
H
12
O
6
/6CO
2
= 0,66.
Tr s E
T
tm ra theo s lng ban ngy trong khong 42 93; tr s bin i
trong ngy trong khong 50 150. Tr s ny nh hn nhiu so vi tr s bit (270
- 380). Bi v thi gian tin hnh tnh bnh qun, phng php o v mi trng o u
khc nhau, s liu trc y phn ln ly t th nghim trong chu, ng nhin khc
vi tr s tm c din tch ln hin nay, hn na s sai khc ny c th c lin quan
ti s nh hng ca nhit di ng.
Quang hp ca cc tng khc nhau trong qun th: thng lng CO
2
, P
z
tin hnh
vi phn cao z c cng thc sau y:
dP
z
d dC
Qp(z) =
dz
= -
dz
(Kz
dz
) (64)
Trong cng thc: Qp(z) l
cng ngun ht CO
2
; Kz l h
s khuch tn dng xoy, C l nng
CO
2
. Qp (z) tng ng vi
cng quang hp thun cc
tng ca qun th. Nu khng dng
Qp (z) m thay vo l thng
lng CO
2
ca cc tng, ly tr s
ny chia cho tng lng quang hp
s tm c tr s ca cc tng so
vi tng quang hp ca qun th.
Hnh 39.2 nu ln mt th d v
o cng quang hp. Hnh ny
cho thy din tch l ca cc tng
(F/Ft) v cng quang hp ca
n v din tch l.
Ngy 26 thng 7, hnh dng ca
F/Ft v P/Pt rt ging nhau, cn
ngy 8 thng 8 th c sai khc tng
i, ni r P/Pt tng di qun
th hu nh khng c quang hp. S
sai khc gia hai ngy c phn

0
1.0
0 0,2 0,4 0,2 0,4 0 1,0
0
0
0,1 0 0,1 0,2 0 1,0
Ft
F
Pt
P
Ft
F
Pt
P
0600-1700
Pt . F
Ft . P


29/7
0600-1700
Pt . F
Ft . P


8/8
1,0
C
h
i

u

c
a
o

t

n
g

i

(
Z
/
H
)

0,5
C
h
i

u

c
a
o

t

n
g

i

(
Z
/
H
)

0,5
Hnh 39.2. Quan h gia cng quang hp
ca cc tng trong qun th ng vi cng
quang hp ca n v din tch l (Inoue, 1968)
F/Ft - din tch l tng i
P/Pt - cng quang hp (ht CO
2
) theo tng
(tr s tng i)
P. Ft/F. Pt = (P/Pt) (F/Ft) - cng quang hp
ca n v din tch l (tr s tng i).
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

nh rt r rng cng quang hp ca n v din tch l.
7. S sinh trng ca qun th cy trng
Sut sinh trng tng i v sut quang hp thun
V mt nh lng ca qun th cy trng, c th biu th bng kch thc ca cy
trng (kch thc hnh hc), cng c th biu th bng khi lng. S sinh trng ca
qun th cy trng th hin chc nng ca h sinh thi ng rung, cho nn dng khi
lng cht kh lm ch tiu l tt nhin. Ni chung, s sinh trng ca sinh vt c dng
hm s m. Blacman l ngi u tin dng quan h phn tch s sinh trng ca
thc vt v ng nu ra cng thc sau y:
1

dW
r =
W

dt


(65)
Trong : r l sut sinh trng (sut sinh trng tng i - vit tt l RGR), tc l
lng tng ca khi lng cht kh W t l vi khi lng cht kh lc . Tnh tch
phn cng thc (65) c:
W = W
0
e
rt
(66)
Trong cng thc, W
0
l khi lng cht kh khi t = 0. Ly logarit hai v, cng thc
(66), thnh cng thc sau:
lnW = l
n
W
0
+ rt (67)
lnW tin hnh t hp theo t s tm c dc r. chnh l nh lut Fourier v
sinh trng thc vt ca Blackman (1919). Kira (1953) theo di s bin i ca tr s r
theo s sinh trng, khng dng cng thc (65) v (66), ng a ra cng thc:
1

dW
W

dt
= r (t) (65a)
W = W
0
exp [
t
0
r (t) dt ] (66 a)
l nh lut Fourier pht trin, dng thch hp cho mi ng cong sinh
trng bt k. tm r bnh qun trong khong t
1
v t
2
, c th dng cng thc sau:
1

dW dlnW lnW lnW
2
- lnW
1
r =
W

dt
=
dt

t
=
t
2
- t
1
(68)
n v ca r l ngy
- 1
, tun
-1
.
Trn thc t sut sinh trng r ca cy trng thng c tr s ln thi k u sinh
trng, sau thp dn. Nh th, do r lin quan vi qu nhiu nhn t nn khng th
tin hnh gii tch. Ngi ta thng dng c quan sn xut ra cht kh, tc l n v
din tch l biu th s tng ca khi lng cht kh v ly lm thc o ca hiu
sut quang hp, gi l sut ng ho NAR, NAR tnh theo cng thc sau:
NAR = 1

dW (69)
45
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

u dt
Trong : u l din tch l. Khi tnh NAR theo tr s o thc c th dng cng
thc sau:
1

dW dlnu dW lnu
NAR =
W

dt
=
du

dt

u

=

lnu
2
- lnu
1
W
2
- W
1
=
u
2
- u
1

t
2
- t
1
(70)
Theo khi nim cu NAR th:
1

dW u 1 dW
r =
W

dt
=
W

u

dt
= LAR LAR (71)
Nh vy, RGR b chia ct thnh NAR v LAR (sut din tch l). LAR l din tch
l ca n v trng lng ca cy trng, theo cng thc (70) cng c th tm c:
u lnW
2
- lnW
1
u
2
- u
1
LAR =
W
=
W
2
- W
1

lnu
2
- lnu
1
(72)
ng thi dng NAR v LAR gii tch s sinh trng ca thc vt s cng t m
hn vi dng ring RGR gii thch.
V s sinh trng ca qun th cy trng, c th biu th khi lng cht kh theo
n v din tch t ai, lng tng ca n gi l sut sinh trng cy trng CGR.
dy 1

dW
CGR =
dt
=
W

dt
F = NAR LAI (73)
Trong , F l din tch l ca n v din tch t ai, gi l ch s din tch l
LAI.
H phng trnh sinh trng:
V m hnh sinh trng cy trng, y xin gii thiu h phng trnh sinh trng
do Rot (1967) xut. Munekata (1967) khi tho lun m hnh chc nng ca nng sut
la nc, nu ln kh nng dng h phng trnh vi phn biu th v mt nh
lng s sinh trng ca cy trng. M hnh ca Rot (1967) ly quan im sau y lm
c s: 1) sinh trng ca cy trng khng phi dng tng trng lng kh ca c th
biu th, m phi ln lt xt theo s sinh trng ca cc c quan; 2) sinh trng ca
cc c quan li chia ra sn phm quang hp mi sn xut lc v sn phm quang hp
c tch tr. Theo quan im th nht, thc vt c chia ra l (1), thn v cnh (2), r
(3) v c quan d tr sinh thc (4). Trng lng cht kh ca cc c quan ny l mf(t).
Theo quan im th hai, gia l v thn va c sn phm quang hp to thnh vi hnh
thc (A
21
- A
12
) v (B
21
- B
12
), ng thi sn phm quang hp c tch tr cng tin
hnh trao i. Ch s vit trong ngoc n 21 l ch s vn chuyn t c quan 2 (thn)
n c quan 1 (l). Cn s vn chuyn quang hp mi t c quan i n c quan j th
biu th bng cng thc sau y:
46
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

47
1 ij
4
1 j
=

=
A
ij
= K
p
P
i

ij
(74)
Trong ,
ij
l t sut vn chuyn sn phm quang hp sn xut ra trong c quan
th i (K
p
P
i
) n trong c quan th j; K
p
l h s tnh i t CO
2
thnh cht kh. Vy c
cng thc sau y:
(75)
V s trao i ca sn phm quang hp c tch tr, n t l vi tng khi lng
cht kh ca cy trng, tc l:
B
IJ
B
JI
=
IJ
. M (76)
Trong ,

=
=
4
mj M
1 j
;
ij
biu th bng hnh thc t s i vi 1 gam tng khi
lng cht kh ca cy trng, l lng trao i sn phm quang hp tch tr gia c
quan th i v th j, do c quan h nh sau:
0 mj
4
1 j , i

=
(77)

ij
= -
ji
(78)
Cn c vo cng thc (74), (75), (76), c th c phng trnh sau y:

= =
+ =
4
1 i
ij j j r
4
1 i
i ij p
M V R K P K
dt
dmj
(79)
Rj trng lng h hp ca c quan th j ; Vj l lng cht kh tng ng, Kp
KR.
ij (t) v ij (t) gi ln lt l ma trn ca sinh trng dinh dng v sinh trng
sinh thc. Nu tm ra ij v ij th vic gii thch sinh trng cy trng s cng c
tin trin hn.
Di y khng dng ij (t), m thay n bng hm s vect bnh qun ca sinh
trng dinh dng
) t ( a
, tc l:

=
=


=
4
1 j
j j R j P
4
1 i
i i R i ij p
) V R K P K (
V R K P K
) t ( a
(80)
Cng hon ton ging nh trn, c th dng cng thc sau y thay ma trn sinh
trng sinh thc bng hm s bnh qun ) t ( b .

=
=
4
1 i
ji
) t ( bj (81)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

48
Theo cng thc (80) v (81) th cng thc (79) c th bin dng nh sau:
dmj dM
dt
= aj (t)
dt
+ bj (t) M (82)
aj (t) trong cng thc (82) c bit theo cng thc (80) l t sut phn phi cht
kh sn xut trong thi gian t v c quan th j. Trong thi gian sinh trng dinh dng,
bj (t) = 0 cho nn:
dmj
aj (t) =
dM
(83)
Trong thi gian sinh trng sinh thc:
dmj
dt
bj (t) =
M
(84)
Hnh 40.2 biu th hai hm s sinh
trng aj v bj ca cc c quan khc nhau
ca i mch o c nc Cng ho
Extnia. Sau khi va mi ny mm, sn
phm quang hp c dng vo sinh
trng ca l v r, t sut ca chng c
chng l 2 :1. Sau khi phn phi xong
vo l v r, s phn phi vo thn cng
bt u tng ln, khi cao nht cn tiu dng
70%, cui cng mi phn phi cho bng.
0ngy 20 40 60 80 160
-8
-4
1964
1965
4
1
2
3
0
4
8
1,0
0,8
0,6
0,4
B
j
x
1
0
3

0,2
0
Thng 5 Thng 6 Thng 7 Thng 8
nh lut phn phi cht kh:
Moritani (1970) tm ra kt qu v
phn phi cht kh sn xut ca nhiu loi
cy trng, nh gia tng khi lng cht kh, khi lng b phn ca la nc v u
tng c quan h nh sau:
Hnh 40.2. Hm s sinh trng ti
cc c quan ca i mch (Rot, 1967)
1. L; 2. R; 3. Thn, 4. Bng
F = W - W
2
(85)
C = (1 - ) W + W
2
(86)
Trong cng thc: F v C tng ng l khi lng ca b phn phin l v khng
phi phin l; W - khi lng ton b; v l hng s kinh nghim.
Cng nh trn, i vi ng v c ci ng tm ra cng thc bc 3 sau y:
F = W - W
2
+ W
3
(85)
C = (1 - ) W + W
2
- W
3
(86)
Cng nh vy, quan h gia khi lng phn trn mt t v r cng c th biu th
bng cng thc bc 3. Trn thc t khi nghin cu vn sinh trng, ni v t sut cu
thnh cc c quan ca cc cy trng khng quan trng bng ni v s phn phi
nhng cht sn xut. Monsi c mt cng thc v sn xut l:
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

W = (p - r) F - r C (87)
Trong : p l lng quang hp; r v r l lng h hp ca F v C. em cng thc
(85) v (86) thay vo cng thc (87):
W = a W - b W
2
(88)
Tin hnh phn tch i vi cng thc (88) (coi a, b l hng s) s c phng
trnh sinh trng sau y:
a/b
W =
1 + Ke
-at
(89)
Trong cng thc: a = (p - r + r); b = (p - r + r); K l hng s tch phn (trn thc
t a, b khng phi l hng s, nn cng thc (89) khng th thnh lp c).
Hon ton ging nh vy, c th t cng thc (85a) v (86a) tm ra:
W = aW - bW
2

+ cW
3
(88a)
KcW
2
2b 2cWb
ln =
cW
2

- bW + a
+

tg
-1

= t (89a)
Trong cng thc, c = (p - r + r)
2
= 4ac - b
2
Cng thc (89) c th l mt ng cong lgarit dng ch S. Moritani (1970)
nghin cu quan h ca v vi nng sut: khi to v nh, th F
max
= (
2
/4), khi
lng kh khi khi lng l t n ln nht (/2) cng ln, trng lng thc thun
ca la nc cng cng ln. Vi u tng, quan h ca nng sut vi , cn cha r.
Ng v ci ng trong trng hp nh v / ln th nng sut cao hn. thi k
u sinh trng, nhn vo quan h:
F W (90)
Trng lng l t l vi tng khi lng cy, cn i vi , v sau ny vn cn
thit phi nghin cu k thm.
8. S cnh tranh ca h sinh thi ng rung
Gia cc thnh phn sinh vt cu thnh h sinh thi c s tc ng ln nhau, l
im cn bn cu thnh h thng. Nhng tc ng ln nhau ny chia ra quan h trong
loi v quan h gia cc loi, chia nh hn th c quan h cnh tranh v quan h h tr.
Quan h gia cc loi coi l cnh tranh th ngoi cnh tranh ra, cn c bt n v sng
gi (k sinh). y ch tho lun vn cnh tranh gia cc loi. Hn na, trong loi
cng c cnh tranh, nu mt qun th tng ln, c th xut hin hin tng gi l
cnh tranh ca hiu qu mt . Di y s ni n vn quan trng nht trong h
sinh thi ng rung l vn cnh tranh gia cc loi, gia cy trng v c di...
C di
C di vn l t dng ch nhng thc vt thn tho nh m con ngi khng
cn, khng ch gii hn ng rung, c nhng thc vt thn tho nh khng c
hoan nghnh trong i sng loi ngi pht sinh trn ng ci, vn hoa, trong rng,
49
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

50
trn thm c u l c di. C di ng rung nng nghip gi l c di ng rung
d phn bit. C di ng rung v th c th hiu l loi t sinh trn t ng rung nh
rung nc, t cn, b rung v t ln cn ngoi cy trng, c nguy hi trc tip hoc
gin tip i vi cy trng. Qun th c di trong ng rung l mt giai on ca din
th. Nu khng c x l ca con ngi th c di mc rm rp, din th v pha thin
nhin pht sinh bin i nhanh chng v hng qun th che ph t cao nht trc y.
Nhng v c canh tc (mt loi hot ng ca con ngi), mi lm cho hng din th
ca qun th c di t hng t nhin quay ngc tr li, dng khi im.
Loi v loi hnh c di ng rung
Phm Hong H (1993) m t 10.500 loi c di Vit Nam bao gm cc loi c
di trn cn v di nc. Nguyn Hng Sn (2000) pht hin 60 loi c di xut
hin trn rung la vng ng bng sng Hng. Theo mc pht sinh, nguy hi ca
c di v phng tr kh d th c th chia ra ba loi: c hi nghim trng, c hi thng
thng v c hi nh. C di nghim trng trn t la c 29 h vi 82 loi c di
(Nguyn Mnh Chnh v Mai Thnh Phng, 2004).
Ni chung thng dng kiu sng lm c trng ca qun th thc vt tin hnh
phn loi. Nh vy, i vi qun th c di cng c th dng kiu sng hay kiu nhn
ging ca chng tin hnh phn loi. Hnh 41.2 biu th s sai khc v s loi theo
kiu sng ca c di cc loi t khc nhau.
C th thy l so vi b ca rung nc v t cn, ni chung thc vt mm mt
t v thc vt mm di t ca ng rung xut hin t hn. t mi khai khn cng
thuc th c di sng lu nm pht sinh cng t. T s thc ca hai mt so snh gia
hai b v trong rung, ngi
chm nom t hn, gn hn vi
trng thi ng c t nhin, cho
nn s loi sng lu nm nhiu.
Th H He G Hy
C di sng
u ma h
C di sng
cui ma h
C di sng
ma ng
R
u

n
g

n

c
B


r
u

n
g

n

t

c

n
B

t

c

n
R
u

n
g

n

c
B


r
u

n
g

n

t

c

n
B

t

c

n
R
u

n
g

n

c
B


r
u

n
g

n

t

c

n
B

t

c

n
0
20
40
60
80
100
%
Arai v ctv. (1951b) xut
pht t lp trng nghin cu
c di ng rung, ra
cch phn loi theo thi k pht
sinh ca c di v theo tnh
thch ng ca chng vi lng
nc trong t (bng 4.2).
T


l
o

i

S pht sinh c di v
lng c di
Hy ly rung nc c
trng v ng lm th d, Arai
v ctv (1951) ch ra mt s
quan h nh hnh 42.2. Qua
quan st pht hin tnh khu vc
Hnh 41.2. Phn loi kiu sng c di ng rung
cc loi t khc nhau v s sai khc v s loi
ca chng (Arai, Yokomori, 1951 b)
Th - thc vt sng 1 nm , H - thc vt mm mt t,
G - thc vt mm di t, He - thc vt sng ng ly,
Hy - thc vt sng nc
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

v thnh phn v lng c di ca qun th c di, c quan h cht ch vi s pht sinh
ca c di, nht l nhng c di ch yu ca rung nc trng v ng.
Nu thi gian t khi rt nc cho la n thi k gieo trng cy v ng ko di,
s lng ht ging A.fulvus [c di sng ma ng (thu)] gim i, s lng ht ging
Stellaria uliginosa [c di sng ma ng (xun)] t b thay i v th m tng ln mt
cch tng i, thm ch dn n bin i thnh phn hp thnh ca qun lc c di (di
ng v pha bn phi theo trc honh ca hnh 42.2). Nu nhit khng kh thi k
gieo trng m mch m thp n di nhit thch hp cho ht c di ny mm th so
snh vi s lng ht ging, t l cy mc mm thp, s lng ht c di cng gim.
Bng 4.2. Phn loi c di ng rung theo thi k pht sinh v
theo tnh thch ng vi lng nc trong t (Arai v ctv. 1951b)
Phn loi theo thi k pht sinh
Phn loi theo tnh thch ng
vi lng nc trong t
C di ma h
Pht sinh t ma xun n
thu
C di a nc
Sng ma h (sm) Pht sinh r thng 4 - 5 a nc (nc )
Sng ma h
(sm, va)
Nh trn, nhng sau
cng nhiu
a nc (m)
Sng ma h (va) Pht sinh r thng 5 - 6 C di a m
Sng ma h
(va, mun)
Pht sinh r thng 5 - 6,
9 - 10
a m (nc, kh)
Sng ma h (mun) Pht sinh r thng 7 - 8 a m (m)
C di ma ng Pht sinh ma thu v xun a m (kh)
Sng ma ng (thu) Pht sinh r thng 9 - 10 C di a hn
Sng ma ng (xun)
Nh trn, nhng ma
xun cng nhiu
a hn (m)
a hn (kh)
Mt khc, trng tip v ng trn rung nc, trc v trng m mch vo ma thu
cng nh trc v cy la nc vo ma xun u phi cy lt t. Nh th cng c th
lm thay i s lng cy ca c di (hnh 42.2). Phn na trn ca hnh l s lng
Alopecurus Fulvus, phn na di l s lng ca Stellaria uliginosa. Cy ma xun
lm ht ging b vi vo trong t, do cch cy khc nhau m s phn b ca ht ging
c cng c khc nhau; nu cy xi nng th ht ging tng mt nhiu, trc khi cy
ma thu, s pht sinh ca hai loi c di u tng.
51
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

52
Alopecurus fulvus trong thi
gian rt nc, s ht ging sng
gim i, khi cy ma thu nu cy
su, t c o lt nhiu, th s
phn b ht ging tng trn tng
ln, s cy ny mm tng ln nhiu.
Trong trng hp cy xi nng, do
ch o t tng mt tng i t
ht ging, cho nn pht sinh c di
li t. Tri li, s phn b ht ging
c Stellaria uliginosa v s cy c
th ny mm nhiu hay t, do
phng thc cy ma xun quyt
nh, sau khi cy ma xun gi
nguyn khng ng n, n khi
cy cng nhiu. Ngc li nu ma
thu tin hnh cy lt su th s cy
mc t hn. Tm li, lng pht
sinh hai loi c ny, do p dng t
hp phng php cy t m c phn ng khc nhau, thnh phn hp thnh qun lc cng
khc nhau. Trong h thng cy lt su ma thu, c Alopecurus fulvus tng ln; cn h
thng cy xi nng ma thu, th t l c Stellaria uliginosa tng ln.
T
h

p

C di ma
ng (thu)
C di
sng cn
C di ma
ng (thu)

C di
sng m
C di ma ng (nc)
C di ma h (c ) n
C di sng cn
C di ma ng (xun)
C di ma h (cn)
C di sng m
S ngy t lc gieo
la n lc gieo mch
Ngn Di


n
h
i

t

k
h

n
g

k
h

c

g
i
e
o

m

c
h

C
a
o

Hnh 42.2. M hnh cu trc c tnh khu vc v
lng c di hnh thnh v thnh phn hp thnh
qun lc c di ca rung nc, c trng tip
v m mch
S cnh tranh gia cy trng v c di
C di cnh tranh nh sng, cht dinh dng, nc vi cy trng ng thi l
ngun truyn bnh v su hi cho cy trng (Trn c Vin, 1998). C di xut hin
trn ng rung s che bt nh sng ca cy, ng thi ht mt phn cht dinh dng
v nc lm gim ph ca t v nh hng n s sinh trng v pht trin ca cy
trng. Trong thc tin sn xut, c di cnh tranh mnh m nht vi la cn vo thi k
28 - 42 ngy sau gieo (Nguyn Mnh Chnh v Mai Thnh Phng, 2004).
Nhiu loi su bnh hi cy trng sinh sng v pht trin c trn c di v n ly
c di lm k ch trung gian t truyn bnh sang cy trng. Theo Nguyn Mnh
Chnh v Mai Thnh Phng (2004), cc loi nm bnh o n, kh vn, su cun l la,
b xt sinh sng c trn nhiu loi c ha bn nh c lng vc, ui phng, c ch.
Kozaka (1965) thng bo c ti 188 loi cy thuc 32 h thc vt l k ch ca nm
trn la v nm tch ra t nhiu cy khc u c kh nng gy bnh cho la Nht Bn.
Vit Nam, bo lc bnh l ni truyn bnh nm bnh kh vn.
V s cnh tranh gia cy trng v c di, ngi ta cn ly cht tit ra ca r lm
mi gii gi l hng nghin cu t gc tc dng ln nhau v hng ho gia thc
vt. Nu kho lo li dng c quan h cnh tranh nh vy gia cy trng v c di, s
c kh nng tr thnh phng php phng tr c di va l tng va chnh xc.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Ngi ta cho rng c hai phng hng phng tr c di. Mt l, nh k thut
canh tc lm bin i mi trng ng rung, to ra iu kin mi trng khng c li
cho s ny mm, n hoa v kt ht ca c di, do m c ch s nhn ging v t
hiu qu phng tr c di. chnh l phng php phng tr bng sinh thi. Hng
th hai l tm cch tr tit c di pht sinh v ang pht sinh, c th gi l phng
php dit tr c di. D phng php no cng u phi nghin cu y sinh l v
sinh thi ca c di, l tng nht l li dng quan h cnh tranh gia cy trng v c
di tin hnh phng tr.
Cnh tranh gia cy trng
Kiu cnh tranh khc loi v cng loi cy trng xy ra rt ph bin trn rung
trng xen, trng gi, rung trng mt loi cy trng. Cnh tranh cng loi l mt nhn
t quan trng to dng ng ca cy v mt cy trng. tng nng sut v hn
ch s cnh tranh cng loi, cc ging cy trng c to ra c l to vi thn thnh gc
nh. hn ch cnh tranh khc loi, ngi ta la chn cc loi cy a sng v a bng
trng xen vi nhau hoc cc cy trng c cc chu k sinh trng hoc kch thc khc
nhau nh xanh trng gi vi ng ng.
Mt khc, cy trng cng nh c di u c kh nng tit ra cc hot cht sinh l
c kh nng km hm s pht trin ca cc loi khc nh mt s loi cy c kh nng
tit ra tinh du.
9. Nng sut ca h sinh thi ng rung
Khi nim v nng sut
Sn xut, vn l thut ng ca kinh t hc, l khi nim ngc li vi tiu dng.
Trn ngha , sn xut trong nng nghip tc l s tc ng vo t nhin v cy
trng thu c nhng vt cht cn thit. V th, ng nhin khng phi ch tt c
cc b phn ca cy trng, m ch l nhng phn cn thit. ng thi cng sinh ra mt
khi nim ngc li vi n, tc l sn xut sinh hc. Gi l sn lng sinh hc trong
mt thi gian nht nh ca mt qun th sinh vt no tc l tng lng cht hu c
m qun th sinh vt sn xut ra trong thi gian . i vi sn lng ca n v
din tch trong thi gian nht nh c gi ring l nng sut. Tng lng nng sut
ca thc vt xanh lc (lng quang hp) gi l sn lng th, sau khi khu tr lng
tiu hao do h hp, phn cn li gi l sn lng thun. trnh ln ln khi nim v
nng sut ca h sinh thi ng rung, cng cn c s phn bit. Khi nu ring sn
lng hoc nng sut, l ch vt cht thu hoch cui cng ca cy trng (tc sn lng),
cn tng sn lng (tc tng sn phm quang hp) th gi l sn lng sinh hc. Ngoi
ra, nng sut quang hp (sn lng quang hp) cn gi l nng sut s cp.
Phng php tnh nng sut sinh hc
Thng thng c hai phng php tnh nng sut sinh hc: (i) cn c vo lng
sinh vt tn ti trn din tch nht nh tc l lng hin cn; (ii) cn c vo s trao i
vt cht. Phng php u li chia ra: 1/ Phng php cn c vo lng hin cn cui
cng hay cao nht c tnh (hiu chnh lng cht kh m ng vt n mt l c th
53
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

tm ra sn lng thun); 2/ Phng php tin hnh tnh lin tc lng hin cn (tin
hnh tnh lp i lp li lng hin cn, tm c nng sut t lng sinh trng gia
cc ln tnh); 3/ Phng php c tnh cn c vo lng hin cn, lng quang hp v
lng h hp (cn c vo lng quang hp v h hp ca n v sinh khi c tnh,
ri xt n lng hin cn cc thi gian tm ra sn lng sinh hc).
Phng php sau l phng php khng ly lng hin cn lm cn c, tc l trc
tip do s trao i ca O
2
v CO
2
v t tm ra s liu, phng php ny c ng
dng rng ri vo vic nghin cu sinh vt ph du, gn y cng bt u ng dng
vo vic nghin cu cy trng.
Nng sut sinh hc ca qun th cy trng
Nng sut sinh hc ca qun th cy trng do s quang hp v h hp ca qun th
quyt nh, nh phn trc nu, c quan h vi s quang hp v h hp ca l n
v cu trc ca qun th. Cng cn phi xt n s h hp ca cc c quan ngoi l. Cc
c im li c tnh di truyn, phng php trng trt v iu kin mi trng ca
cy trng quyt nh. Nng sut sinh hc ln nht mt ngy ca cc cy trng khc
nhau c nu r trong bng 5.2, ln nht l hng dng v ng:0,79 v 0,52 tn/ha
tng i nh l la nc v ci ng: 0,3 tn/ha. Nu i thnh tr s ca mt ngy,
c th dng tr s bnh qun ca thi gian sinh trng biu th: 0,05 - 0,15. Cc tr s
ny so vi cc iu kin trng trt khc, th d nui trong dung dch, th tr s trong
thi gian rt ngn l 1,22. iu kin ca h sinh thi t nhin th gn vi cy trng,
hoc hi thp hn.
Hy ly sn lng thun v tng sn lng trong c thi gian sinh trng hoc mt
nm (cy lu nm) ca cc loi cy trng so snh vi h sinh thi t nhin.
Bng 5.2. Nng sut sinh hc ca cc cy trng khc nhau (Willson, 1968)
Nng sut sinh hc mt ngy (tr s ln nht) tn/ha. ngy
Hng dng 0,79
Ng 0,52
C xu ng 0,51
Ma 0,38
Ci c 0,34
La nc 0,33
Ci ng 0,31
Nui cy nhn to n bo 1,22
To 0,86
Tr s bnh qun ca cy trng trong thi gian sinh trng 0,05 - 0,15
H sinh thi t nhin
Hng b Typha latifolia 0,53
Sa tho Scirpus lacustris 0,28
Pteridium aquilimum 0,20
54
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Bng 5.2 cho thy, trong s cc cy trng, tr ma v ng ra, sn lng thun u
l kt qu tch cc trng trt vi mc tiu l nng sut cao, t n con s rt cao. So
vi tnh hnh , tr s ca h sinh thi rng v ng c l gn ging nhau, thng u
thp hn cy trng. Cn v tng sn lng, h sinh thi rng th hin r tr s rt cao,
trong h sinh thi ng rung, th d la nc, i mch gn bng hoc hi cao hn
rng. Mt khc, cc thi gian sinh trng khc nhau, nu dng tr s mt ngy biu
th, th sn lng thun hay tng sn lng ca h sinh thi ng rung u biu th tr
s kh cao, nhng ng c ch l rng cy Zelcova serrata (Mark), loi rng l
rng rng l cng c tr s gn ging nh th.
Nng sut v sn lng sinh hc
Ni chung, cy trng ly ht v cy trng ly thn l, c cu trc quyt nh sn
lng rt khng ging nhau. V cy trng ly ht phi tri qua hai qu trnh: qu trnh
hnh thnh c quan vt cht thu hoch v qu trnh tch lu sn phm quang hp vo
trong c quan . Vi cy trng ly thn l th ch yu l qu trnh hnh thnh c quan.
Th d la nc, quan h gia sn lng sinh hc (nng sut cht kh) v sn
lng, c th ly thi k tr bng lm ranh gii, trong khong thi gian mt thng trc
v sau tr bng l khng ging nhau. Nng sut cht kh trc khi tr nhiu hay t c
quan h cht ch vi s ht th tinh (s hoa) nhiu hay t, ch cn nh c s hoa, mc
d nng sut cht kh sau khi tr vn tng nhiu ln, nhng sau khi sn lng t n
mc nht nh th khng tng tng ng na. Ngoi ra, gia nng sut cht kh sau
khi tr v khi lng ht chc cng c tng quan cht ch, tc l nng sut cht kh
cng cao, khi lng nghn ht cng cng ln. Tm li nng sut cht kh v hai qu
trnh quyt nh sn lng c mi lin h ring bit, nn mi hnh thnh kt qu ni
trn. C th thy quan h gia nng sut v sn lng sinh hc kh phc tp.
Tnh khu vc ca nng sut
Nng sut la nc ca Vit Nam, theo s liu ca Tng cc Thng k (2004) v
B Nng nghip v PTNT, bnh qun cc nm 2000-2003 dao ng t 4,24-4,63 tn/ha,
tnh Thi Bnh t nng sut cao nht 6,3 tn/ha; tnh Lai Chu thp nht vi 2,54 tn/ha
(2002).
Ngi ta cho rng c th dng hm s lng chiu sng mt tri (thi gian chiu
sng ca mt tri) v nhit khng kh ca thi k chn biu th s sai khc v khu
vc. Murata gi l s ch nng sut kh tng, thu c bng cng thc sau y:
Sf(t) = S [1,20 - 0,021 (t - 21,5)
2
] (93)
Trong : S l lng chiu sng bnh qun ngy t trc khi tr 10 ngy n sau
khi tr 30 ngy, t nhit khng kh bnh qun ngy trong thi gian trn, Sf (t) l s ch
nng sut kh tng, f(t) l ch s nng sut nhit. f(t) trong cng thc (93) tm c
bng phng trnh bc 2, c ngha l c tn ti nhit ti thch. Habu ch ra rng:
YR = S [4,14 - 0,13 (21,4 - R)
2
] (94)
Trong : YR l s ch lng chn kh hu, S l tng s gi chiu sng trong 40
ngy sau khi tr, R l nhit khng kh bnh qun trong thi gian ni trn.
55
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

T tm ra thi k tr bng cho tr s YR tr thnh cc i ca cc khu vc,
nh vy nu r quan h phn b a l ca tr s cc i ca YR. Nhng kt qu ny
ni rt r tnh khu vc ca nng sut la nc, ng nhin cn phi xt n quan h
v s sai khc ca nng sut ca t vi s hnh thnh tnh khu vc ca nng sut tc
ng. Hn na, trong s hng f(t) ca cng thc (93) cng phi xt c n tc ng ca
con ngi nh k thut trng trt.
Bng 6.2. Sn lng thun v tng sn lng ca cc h sinh thi
Trong sut thi gian
sinh trng
P
n
P
n
ngy
P
g

ngy
Cy trng
hoc thc
vt u th
ca h sinh
thi
a im
Sn
lng
thun
P
n
Tng
sn
lng
P
g
Lng
h hp
R
P
g
tn/ha/ngy
Nng
sut
tn/ha
ng rung
Ma 33,0 0,10
La m Chiba 29,0* 52,7* 32,7* 0,55 0,13 0,24 10,01
La nc Akita 27,8* 52,5* 24,7* 0,53 0,19 0,37 10,51
Ng Nagano 26,5 0,21
u tng Miyagi 20,9* 29,2* 18,3* 0,53 0,17 0,32 7,85
Ci ng Hokaio 17,1 0,10 11,40
La nc Nagatama 16,3 30,8* 14,5* 0,53 0,14 0,26 6,24
i mch Kitama 15,3 26,4* 11,1* 0,58 0,08 0,13 5,20
u tng Iwate 8,0 15,0* 7,0* 0,53 0,07 0,13 2,96
Rng
Rng ma
nhit i
Thi Lan 28,6 123,2 94,6 0,23 0,08 0,34
Rng trng
chu u
an Mch 13,5 23,5 10,0 0,57 0,07 0,12
Rng trng
samu
Kumamoto *** 84,1 55,0 0,35 0,09 0,25
- nt - Oita *** 52,6 36,4 0,31 0,07 0,22
ng c lu
nm
Omuri
28,6
**

Csium
Nipponirum
Gunma
20,6
**

** Tr s ln nht ca lng hin cn
56
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

57
10. M hnh ho h thng h sinh thi ng rung
Gn y, phng php k thut hc h thng nu r chc nng h sinh thi ng
rung, ngy cng c ng dng rng ri trong lnh vc sinh thi hc. Chi tit s c
ni n chng sau. y ch thuyt minh m hnh ho ca cc h thng ph
(subsystem) cu thnh h sinh thi ng rung.
M hnh ho mi trng CO
2
trong qun th cy trng
iu kin CO
2
trong v ngoi qun th cy trng c quan h cht ch vi quang
hp ca bn thn cy trng, iu kin khuch tn dng xoy ca tng kh chung quanh,
lng h hp ca t... Do , nu r iu kin mi trng CO
2
s gip ta tm hiu
quang hp ca qun th cy trng.
M hnh khuch tn: Trong qun th cy trng, khuch tn CO
2
vi mc bt k
c biu hin bng cng thc sau:
d dC
-
dz
(K
dz
) = - f
L
(z) p (z) + f
L
(z) r
p
(95)
Trong : C l nng CO
2
(mgCO
2
/cm
3
), K: h s khuch tn dng xoy (cm
2
/s),
f
L
(z): mt din tch l (cm
2
/cm
3
), p (z): cng ht CO
2
(quang hp) ca n v
din tch l (g CO
2
/cm
2
.s), r
p
: cng thi CO
2
(h hp) ca n v din tch l (g
CO
2
/cm
2
.s).
Bit c s bin i trong qun th cy trng ca h s khuch tn, mt din
tch l, cng quang hp v h hp, tnh tch phn i vi cng thc (95) l c th
tm c s bin i theo chiu cao ca thng lng CO
2
v phn b CO
2
trong qun
th. V cng quang hp, cng h hp, h s khuch tn v mc lt qua ca
nh sng c th dng cc cng thc sau:
Q
P = D
c
max C
a + Q
(96)
r
p
= hng s = p

(97)
K = K
H
- (H - z) (98)


=

H
z
L H
' dz ) ' z ( f k exp Q
1
K
Q

(99)
Trong cng thc trn, D
c
max l tc trao i iu kin bo ho nh sng, Q:
cng bc x mt tri, P

= D
c
, max. C
a
cng quang hp iu kin bo ho nh
sng, H: cao ca qun th, v l hng s, k l h s tiu tn, l mc (t l) nh
sng lt qua ca l.
By gi chng ta hy thng qua K ni r cu trc hnh hc ca qun th. Tnh
tch phn theo z i vi cng thc (95) bin i ca cc s lng c lin quan l thnh
ng thng th lp mng h kp gia hai cao gn nhau c quan h gn ng nh sau:
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

58
(C
j
+ 1 + C
j
) p
j
1 j , j C +
=
2
1 j , j C + -
K
j
. h (100)
1
P
j +1
(r
p
. F + p
j
) - D
c, max
, F
a + Q
1 j , j C + (101)
P
j
v K
j
l thng lng CO
2
v h s khuch tn mc j; F: s gia din tch l trong
tng, Q: tr s bnh qun cng nh sng mt l trong mc j v j + 1. Cc thng s ly
cc tr s sau y: D
c,max
= 0,41; a = 0,73; H = 200 cm; = 0,063; = 0,1; h = 10 cm; K
H
=
125, 250; 500; 1000; 2000; 3000 cm
2
/s; k = 0,3; 0,5; 0,75; 1,0; F
t
= 1,0; 2,0; 4,0; 6,0; 8,0;
10,0; Q
H
: 0,0; 0,1; 0,2; 0,4; 0,6; 0,8; 1,0; 1,5 ly/min; P
S
: 0,0; 2,0; 4,0; (10
-8
g CO
2
/cm
2
.s)
Ft l ch s din tch, P
S
l cng thi CO
2
ca mt t. Theo cc t hp ny,
tnh ton khong 25000 th d, nghin cu s bin i theo chiu cao ca thng lng
v phn b CO
2
trong qun th m hnh, mt phn kt qu th hin hnh 43.2 v 44.2.
Quan h cu trc hnh hc ca qun th cy trng v s phn b nng CO
2
:
Mt ct phn b ca CO
2
c quan h cht ch vi rm rp ca qun th, nng CO
2

h thp theo mt din tch l tng ln, cui cng hnh thnh mt ct phn b m nng
gia qun th l thp nht (hnh 43.2). im ny ni r c s b sung CO
2
t mt
t v tng khng kh gn mt t. Trong qun th m h s tiu tn tng i ln (tc
loi hnh l nm ngang), xu th ny rt r rng. Trong trng hp h s khuch tn ln
(tc gi ln), nng CO
2
chnh lch nh. Nu mt din tch l ln, th s chnh
lch nng CO
2
do h s khuch tn ln hay nh gy ra, tng trn v tng di ca
qun th s l ngc li.

















Hnh 43.2. Quan h ca nng CO
2
vi din tch l, h s tiu tn, h s khuch tn,
trong qun th m hnh (Uchijima v Inuoe, 1970).
Ghi ch: ng lin biu th trng hp t c CO
2
thi ra, ng chm chm biu th khng c
CO
2
thi ra Lng chiu sng 1,0 ly/min.
H s khuch tn c 5 on bin i, h s tiu tn c 4 on bin i
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

59
Trng hp mt t khng thi ra CO
2
(hnh 43.2, ng chm chm), ch s din
tch l 2 tr xung, khng c quan h vi k, nng CO
2
gim xung theo su. Khi
ch s din tch l t 8 tr ln, nu k ln, th xut hin mt ct phn b CO
2
r rt. l
v l tng di do nh sng lt qua khng tt, do tng ln h hp thi ra CO
2
. Mt
khc, nng CO
2
bnh qun trong qun th gim xung theo s tng ln ca ch s
din tch l, khi n mt ch s no th tr nn thp nht, nu vt qu ch s , nh
thy r trong hnh, th li tng ln. Khi k = 0,3; 0,5; Ft = 4,0; khi k = 0,7; 1,0; Ft = 2,0.
















Hnh 44.2. S bin i thng lng CO
2
trong qun th m hnh do mt din tch l,
h s tiu tn v lng chiu sng gy ra (Uchijima v Inoue, 1970)
H s khuch tn dng xoy tnh ton theo K
H
= 2.000cm
2
/s, Ps = 2 x 10
-8
gCO
2
/cm
2
.s

10
20
30
0 4 8 12
Ch s din tch l

0
Q
H
=1,5 ly/min K
H
= 3000
cm
2
/s
k
0,75
1,0
0,3
0,5







Q
u
a
n
g

h

p

t
h
u

n

c

a

q
u

n

t
h


k
=
1
0
3
g

C
O
2
/
c
m
3
.
S

0,3
0,5
0,75
1,0
k
K
H
= 125
cm
2
/s
20
10
0
0 4 8 12
Ch s din tch l
Q
H
=1,5 ly/min



Hnh 45.2. Quan h ca ch s din tch l vi cng quang hp thun
(Uchijima v Inoue, 1970)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

60
Thng lng CO
2
v cu trc hnh hc ca qun th: Thng lng CO
2
trong
trng hp l nm ngang (k = 1,0) v l thng ng (k = 0,3) nu r nh hnh 44.2.
ng cong ca thng lng v theo 2,0 x 10
-8
CO
2
/cm
2
s di ng v bn phi. Loi
hnh l thng ng (k = 0,3), khi Ft = 1,0, thng lng bin i chm, nu Ft tng ln
ln, th thng lng CO
2
h thp rt nhanh, khi n mt su no , dng kh t
hng i xung quay li thnh hng i ln (phn v ng ngang trong hnh). su
lm cho dng kh i hng ngc li l ng thun vi tng nng thp nht ca
hnh 44.2. Ngoi ra, Ft nu t n mt mc no tr ln th s xut hin tng ln
nht dng kh hng ln trn. Cng chiu sng ca tng ny gi l im b sng.
T tng ny tr ln, tc l tng nng CO
2
thp nht tr xung, CO
2
ht trong tng
ny l do mt t v tng di h hp thi ra. Hnh 45.2 cn ni r mt im quan trng
l: c mt ch s din tch l m y cng quang hp thun ca qun th ln nht.
gi l din tch l tt nht, tr s ny l 8 khi k = 0,3 v l 4 khi k = 1,0, m sut
quang hp thun li gn nh khng i. Trong trng hp K
H
= 3.000, k = 0,3, cng
quang hp thun ln nht ca qun th l 22,1, khi k = 0,5 l 22,2; khi k = 0,75 l
21,3; khi k = 1,0 l 19,9 (u 10
-8
g CO
2
/cm
2
.s).
Nu K
H
= 125, tc l trong trng hp tc gi ht sc yu, K = 0,3, sut quang
hp thun ln nht ca qun th l 14,7; khi k = 0,5 l 15,0; khi k = 0,75 l 14,7; khi k =
1,0 l 14,1; ngha l gim kh nhiu so vi trng hp K
H
= 3.000. C th thy r l
quang hp chu nh hng ca h s khuch tn, tc l ch gi. Do , iu
kin chiu sng mnh, h s khuch tn c nh hng rt mnh ti thng lng CO
2

(quang hp ca qun th), nhng trng hp chiu sng yu, tc l Q
H
0,4 erg/min, th
hu nh khng c s khc bit.
WST
WLV
RES
WRT C
PHR
PGRW RGRR
GRW
GLW
(TS)
RSP
GST
T nhng iu nu trn
cho thy mt ct phn b nng
CO
2
trong qun th chu nh
hng rt mnh ca s thi CO
2

t mt t, chiu sng, lt
qua ca nh sng v h s
khuch tn, ng nhin cu
trc ca qun th cng c nh
hng mc nht nh.
M hnh ho sinh trng
ca qun th cy trng: Wit
lp m hnh c bn ca s sinh
trng cy trng (gi l
Elementary Crop growth
Simulator - ELCROS). u tin
l kho tn tr sn phm quang
hp (RES). Hp cht gluxit ca
Hnh 46.2. Kiu m hnh c bn ca s sinh trng
ca qun th cy trng (Wit, Brower, De Vries 1
PHR- qu
970)
ang hp, RSP- h hp, RES- kho d tr, GST- sinh
n
trng khi lng thn, GLW- sinh trng khi lng l,
GRW- sinh trng khi lng r, WRT- khi lng c quan
r, WST- Khi lng c quan thn, WLV- Khi lng c qua
l, PGRW- Lng sinh trng c kh nng ca khi lng r,
RGRR- Sut sinh trng tng i ca r, TS- nhit t
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

sn phm quang hp, ng thi do h hp, dng mt mt s hp cht gluxit. Hp
cht gluxit tiu dng cho s tng thm trng lng ca r, l v thn, cng t kho tn
tr m ra. l WRT, WLV v WST biu th 4 gc ca hnh 46.2. S sinh trng ca
r tin hnh theo phng thc nu pha di hnh. Nu cho trc nhit t, , th c
th cn c vo ng cong pha di bn phi hnh tm ra sut sinh trng tng i
ca r (RGRR). ng cong ny khc nhau do loi cy trng, RGRR v s sinh trng
khng c quan h trc tip, nhng tch ca RGRR v khi lng m non C lin quan ti
s sinh trng trong khi lng c quan r, tc l lng sinh trng c kh nng ca r
PGRW pht sinh quan h vi s sinh trng. Trong kho d tr cn di do, PGRW s
tr thnh lng sinh trng khi lng ca r thc t (GRW). ng v lin trong hnh
biu th s lu chuyn ca cht hu c (kg CH
2
O/ha. ngy), ng chm chm biu th
s lu chuyn thng tin.
Trong tnh ton thc t, theo thi gian tnh ton tr s bnh qun ca 1 gi. By
gi ly cy ng lm i tng gi thit lp m hnh theo trnh t nh hnh 46.2.
Khi tnh ton cng quang hp, c th khng xt tui nm ca l, ly hm s sin
ca din tch l v cao mt tri lp bng, n l s liu trong trnh t. nhit l
gi thit ging nh nhit khng kh (cch mt t 10 cm tr ln), nhit khng kh
2 gi, bin dao ng nh l 0,45 ln.
Sinh trng ca l tm c theo phng thc sau y. Khi nh mm l 0, thi k
hoa c xut hin l 1 DVS bc pht trin (development stage) cn c vo tui mi
ngy (quyt nh nhit v di chiu sng) tm ra sut pht trin (development
rate - DVR). Li cn c vo s phn b DVS ca l quyt nh s phn b ca tui
(age). Lng sinh trng ca l (GLG) l tch ca sut sinh trng tng i c
quyt nh do nhit khng kh v lng sinh trng c kh nng ca l (WLVC) tm
c t phn b tui, cng tc l cn c vo lng sinh trng c kh nng ca l
(PGLV) v t sut tn tr (RPR) tm ra. Ngoi ra, t sut sinh trng ca trng
lng l (GLW) c th tm ra lng sinh trng ca din tch l (GLA).
H hp c th chia ra lm h hp tiu dng cho sinh trng v h hp tiu dng
cho duy tr c th thc vt. H hp cho sinh trng tm c bng cch ly lng sinh
trng ca r thn l nhn vi 0,33. H hp cho duy tr tm c bng cch ly tng s
khi lng kh ca l cc tui khc nhau c xt n c hm lng m protein vi h
s 0,08.
Ly mt th d c th biu th nh hnh 47.2, nu r s bin i trong ngy. Ban
m, t sut tn tr (RPR) thp xung, ban ngy li tng ln. Tng ng nh th, l v
r cng c sinh trng. n trc lc mt tri ln th xut hin nh cao sinh trng,
l do s khi phc p lc gin n c lin quan ti s gim thp bc hi m c. Sau khi
mt tri ln, sinh trng lp tc gim xung. Mt khc, phn lin h gia h hp vi
lng sinh trng cng rt ln, s tng ln ca h hp bt u chm hn so vi quang
hp. Hot ng quang hp c tng ln cht t vo bui tra, l do din tch l tng
ln m c.
61
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

62
thun.

0
300
200
100
29-1-1969
4
2
0
0 4 8 12 16 20 24 gi
LEAF AREA
= 1,3 ha/ha
Lng sinh trng ca r
Lng sinh trng ca l
kg
ha ngy
Tng khi lng =1146 kg/ha
Quang hp



H hp
Ng 18 ngy
Quang hp 1 ngy = 272kg/ha
H hp 1 ngy = 35kg/ha
kg CH
2
0
Ngy
300
200
100
400
Cal
cm
3
gi
3
2
0
mm
Ngy
Quang

H hp
TA = 20
0
C
TS = 20
0
C
REL.HUM = 65%



















Hnh 47.2. M hnh ho s sinh trng ca ng
( Wit v ctv, 1970)

kg mm
ha
15.000
5.000
10.000
Tr s o c
Tr s tnh ton
0 20 50 75 100
S ngy ny mm
Hnh 48.2. i chiu o thc v m hnh
sinh trng ca ng ( Wit v ctv, 1970)
3
3
1
1: California
2: Iowa
3: H Lan
2
2
1
Hnh 48.2 l s i chiu gia kt
qu ca m hnh ho v tr s o thc.
l s so snh tr s tnh ton da
vo v ngy ny mm v s hiu kh
tng cc a im thuc California,
Iowa, H Lan vi tr s o thc cy
ng. ng lin v ng chm chm
trong hnh tng ng biu th tr s o
thc v tr s tnh ton, ta thy c hai
tr s khc nhau rt ln. Nhng nu
cn c vo dc ca ng cong
trong hnh tm ra sut sinh trng
ca cy trng CGR (crop growth rate)
th d im no, hai tr s u rt ng
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Wit v ctv (1970) cho rng iu ny ni ln kh nng d bo ca m hnh
ELC
n th, t v khng kh khng c trao i nhit
v n
i dng lng chy vo lng r.
o vch mng ca l,

t bo khuch tn ln b mt l.
ng kh gn .
ROS l ng tin cy. ng nhin, vi mc nh vy l rt khng y . S tm
ti bn thn m hnh c nhin l quan trng, vic nghin cu cc thng s trong trnh t
cng rt quan trng. V vy, khng nhng cn nghin cu li cc s liu c m nht
nh cn cn tin hnh thc nghim ph hp vi mc ch.
M hnh ho s thot hi nc
Trc ht gi thit gia t v qu
c. i vi qun th pht trin y th gi thit ny l gn ng. C th chia s
thot hi nc thnh 4 qu trnh:
1. Qu trnh nc trong t v
2. Qu trnh nc dng lng t b mt lng r chy v t b
b mt ny bin thnh hi nc.
3. Qu trnh hi nc thng qua khe
4. Qu trnh trao i hi nc gia khng kh tip xc b mt l vi kh
Di y s ln lt nghin cu cc qu trnh
S vn ng nc t t ln lng r
Trong h thng ph t t ln lng r ny, gi thit s vn ng ca nc trng
thi
Q
1
= 2 l
2

*
k (102)
nht nh, c th biu th nh sau:
dW
dl
Trong cng thc trn, phn h ca lng r l khi bn cu bn knh l
o
, gi thit
mt
(103)
ch b
mt
g t h g v lng r ong
n
p th
nc ng mc trong t l khi bn cu bn knh l, lng nc chy vo trong
bn cu l Q
1
,
*
l mt (trng khi) ca nc, W l biu th nc trong t, tin
hnh tch phn cng thc (102), cho l ln hn nhiu so vi l
0
, th:
Q
1
= 2
*
kl
0
(W - W
R
)
W v W
R
tng ng l nc trong t cch lng r kh xa v ngay
lng r. W c th coi l bng nc bnh qun ca t, ly s lng r j (1/cm
3
)
trong dung tch t tu nhn vi hai v ca cng thc trn, ta c cng thc sau:
Q
*
= j

Q
1
= 2 j
*
kl
0
(w - w
R
) (104)
Q
*
biu th thng lng nc bnh qun tron n phn b tr
v dung tch t W. Cho nn ly dy tng t vng r zk nhn vi hai v ca
cng thc (104) l c th tm c tng thng lng nc i vi ton b lng r trn
n v din tch ng rung trong n v thi gian.
Q
R
= ks (W - W
R
)
63
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Trong : Q
R
= Q
*
z
K
(g/cm
2
.s) (105)
bo nhu m l
ks = 2 j
*
l
0
kz
k
S vn ng nc t lng r n t
ra, biu th nh cng thc sau: S vn ng nc y ch yu do lc hp dn gy
P
S
Q = (106)
g
z
Trong vt thi ra (g/cm
2
.s), l h s truyn d a
cc
(107)
, Q l n c thc n nc c
m khc nhau ca thc vt, g l gia tc trng trng, P
S
/
z
l dc ca sc ht
(bar/cm). Do s vn ng ca nc trong c th thc vt khng phi l nht nh, nn
i vi mt n v din tch ng rung, cng thc (106) phi i thnh:
Q V

=
*
z t

Trong cng th n, thi gian, V - hm lng nc trong n v dung tch ca
c th
(108)
c tr t l
thc vt, t tm ra c cng thc sau:
P
S
V
( . ) =
*
z g z

t
Trong , n v u, a sc ht P
S
, p sut thm thu lc
dn
(109)
P

1
V ( +
1
) v
P
S
v V c li h i nha gi P
0
, p
n P
T
c quan h nh sau:
P
S
= P
0
- P
T

(110) P
S
= 2P
00
+
0
V
0
Trong k u phn ly cht nguyn sinh, V
ban
(111)
, P

- p sut thm thu hi bt


0
- th tch
u ca t bo,
1
- h s ca ng thng quan h gia th tch t bo (V - V
0
) v
p lc dn n. Cng thc (110) nu xt n quan h ca p lc thm thu v p lc dn
n, th tch t bo vi dung dch lm long, th tm c:
P
S
P

V
= - ( -
1
)
z

V
0 t
Ly cng c (108) thay vo cng thc (111):
P
S
P

V
0
) (112)
th
P
S
V
0
( ) = -
*
(
z g
z
+
1
t
s , l hm ca
n, nh P
S
/t = 0, gi thit n l mt trng thi
nht
h s z.
cng thc (112) n gin h
nh. Tin hnh tch phn t z = 0 n z = z
H
s c cng thc sau:
( )
R . S 1 . S
H
O
P
P P
dz
g
1
Q


(113)
64
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Trong : P
S.1
l lc hp dn ca l, P
S.R
l lc hp dn ca r, t phn trc du
ngoc v phi ca cng thc (113) l , th:
Q
F
= (P
S.1
- P
S.R
) (114)
cng thc (105) v (114) vi nhau s c
cng l l.
c
l tc trao i (cm/s)
trng thi nht nh nh vy, kt hp
thc biu th s vn ng nc t t n
S vn ng n t t ln l
Gia p lc ca nc trong t v nc trong t c quan h nh sau:

2
Q
G.R
= P
G.O
(
W
R
+
2
) (115)
n
Trong : Q
G.R
v P
G.O
tng ng l p lc ca nc trong t khi nc trong t
l W
R
v W
R
= 0;
2
v n l l ng s kinh nghim.
Kh W
R
t th t c
h
cng thc (105) v (115) m cng thc sau:


P
R . G

= 1
P
W k Q
n
1
O . G
2 S


R
(116)
b mt lng r P
G.O
= P
S.R
, do quan h ca thng lng dng nc trong n
v din tch ng rung chy v t bo nhu m c l vi nc trong t t
r, sc ht ca l nh sau:
a ng t b
QP P

1 Q
P

P
S.L
=

+ (117)
[

2
(W -
K
S
) + 1] n
Q
P
v Q
R
rng thi nht nh l bng nhau. Theo kt qu thc nghim n = 3, P

=
25 bar, = 0,063; nh vy tm ra quan h Q
P
= f(P
S.L
,W) biu th nh ng
t nu nc trong t g ln, t v tr h ng thng.
t
hnh 49.2, ch
tn h Q
P
P
S.L
thnh quan
S vn ng nc t t bo nhu m l n b mt l
Nc t dng lng bin thnh dng hi, thng pht sinh b mt l, nhng cng
c th b mt t bo nhu m. S trao i nc gia l v b mt l ch yu l thng
qua t hi nc) gi l thot hi
cutin, phn ny ch c 5 - 10% tng lng thot hi, nu b qua khng tnh th c
cng
z
kh khng. Nc t ngoi khe kh khng thi ra (tho
thc sau y:
Am
E
T
=
a
(q
o
- q
s
) (118)
65
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

66
t hi nc (g/cm
2
.s),

a
- mt khng kh, A
m

khuch tn phn t, on cc
t b mt t bo nhu m n b t l,
q
o
v t l m tng i ca
khn
nh ca D
I
.
Trong :
E
T
- lng tho
- h s
z l kh g
m
h
q
s
ln l
g kh b mt t bo nhu m ca l
v b mt l.
t D
I
= A
m
/z th A
m
= 0,26 cm
2
/s,
z - ln lt l dy ca l v mt na
ca khong cch gia kh khng tc l
bng 60 - 600 . Vy D
I
= 13 - 48 cm/s,
l tr s c t
Vn ng nc t b mt l n
khng kh
Trao i nc gia b mt l vi khn
E = D
1
3
4 5 6
7
0 4 8 12 16 20 24
P
SL
bar
. Quan h ca nc vn chuyn
l bo nhu m ca l v sc ht ca l,
nc trong t hu hiu
W - Nc trong t h % trung bnh)
0,8
Q
P

m
m
/
g
i
8 14
1,0
0,6


2
0,4
0,2

Hnh 49.2

0

n t
Q
P
- Lng dng nc, P
S.L
- Sc ht ca l
u hiu (s
g
a S 2
ng
m
118) v (119) th:
(q
o
q
kh l:
(119)
T a
( Q - q )
Trong cng thc, D
a
- tc trao i gi
ng l m tng i ca khng kh b
cng thc (
a b mt l vi khng kh, q
S
v q
2
t
t l v ch cao 2cm. Kh mt q
S
t
D D
1 a
E
T
=
a
D
1
+ D
a
a
) (120)
Coi m tng i ca khng kh b mt t bo nhu m l bng m tng
i bo ho ca nhit l, khai trin cng thc Taylo i vi q
O
, c cng thc sau:
C
p
D
a
E
T
=
1
+
+ ( D
a
d +
C
P
R)
D
C
p
C
p
+ L
a
(121)
Trong cng thc, C
P
l t nhit ca khng kh, L - tim nhit ho kh, - vi h s
ca ng cong m bo ho nhit.
By gi it: gi th
C
p
P
a
b
1
=
C
p
+ L

b
2
=
Cp + L
D
a
C
p
b
3
=
Cp + L
(122)

Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

67
Cng thc (121) c th tr :
1

E
T
=
b
thnh
3

1 +
D
(b D
a
d + b
2
R) (123)
1
L
Nng lng bc hi nc:
b
1
D
a
d + R
Do c cng th u:

3
E
p
= b
2
(124)
c sa
E
T
1
=
E
P
b
D
L
(125)
1 +
Khi D
L
nh, nu b
3
ln, th E
T
/ E
P
= 0,96. Nu b
3
, D
L
l tr s trung gian, th E
T
/ E
P

= 0,99, rt gn v 1. D c quyt nh m ca kh khng, cho nn:
D
L
= f ( P
T
(126)
Theo cng thc (100) tm c:
L
P
OO
+
1
V
O
(127)
i
L

)

1
V
O
( P
OO
- P
S. L
)
D = f

Trong cng thc, P


OO
l V
O
ca t bo iu tit kh khng, xc nh dng hm s
ca cng thc (127), cn c kiu m hnh ton hc ca vn ng kh khng.
Da vo c (1 ng thc 25) v (127), tm ra:

=
3 T
OO T
OO 3
T P
T 3
)] 1
k
E
W (
1
[
P E
' P f
E E
E b

S 2
Hnh 50.2 biu th E
T
= f (W, P
S.L
) v E
T
= f (E
P
, P
S.L
) . Tung ca giao im bt
k ca ng cong hm s sau v ng cong hm s trc khi W = hng s, biu th
lng bc hi vi tr s nng lng bc hi khc nhau.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

S thot hi nc v nng lng bc hi nc


i dng kt qu P
.L.S
= f(E
P
), ng thi cn
nu r E
T
= f (W) khi E
P
= const v W). Nh vy tm ra quan h
ca ng lng bc hi nc trong t v thot hi nc (hnh 50.2) khi tr s W nh,
E
T
= f (W) m
a qun
tnh ton theo s liu kh tng l bng nhau,
tc l:

1
2
5
6 7 8
14
3
4












L ni trn, dn quan h E
T
= f (E
P
) v
c quan h P
S.L
= f (
n
tr thnh ng thng, khi tr s W ln th E
T
/ E
P
bng 1. im
ng thng pht sinh cong tc l hm lng nc trong t ti hn (W
O
).
T rt ra kt lun nh sau: t m t y , lng thot hi nc c
th rm rp v nng lng bc hi nc
E
T
= E
P
(129)
Quan h ny c thnh lp trong iu kin W > W
C
, W
C
biu th nh sau:
W
C
= W
C.min
+
5
E
P
(130)
Trong :
E
T
2
(P
O O
)
1/3

B
=
k
S

3(P
S.L
)
4/3

W
C.min
l hm lng nc trong t ti hn khi E
P
= 0, biu th nh sau:
P
O O
)
1/3
W = [(
C.min 2
P
O O
- P
T
-1] (131)
0 4 8 2 24
Sc ht n ca l (P
L
)
1 16 20
c
10
E
T
=f(W
T
.P
L
)
E
T
=f(E
P
.P
L
)
0
1,0
1,2

n
g

c

(
m
m
/
g
i

)

Hnh 50. u ca cng bc hi v c ht nc ca l
t Q ( ); E
T
= f(E
P
,P
L
), tin hnh tnh ton theo Qp = E
T
,
ch s ng v lin l % nc trong t, trn ng chm chm
biu th n ng bc hi
0,6
0,8
c

h

i

n

0,4


b
0,2
C
2. Q an h s

P
= f
trn
n
w,P
L

g l
68
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

69
c c,
nc tc trao i v nhit khng kh. Quan h ny im ho tr
xung mi c thnh lp. Nu n t thiu hn na, th cn phi nghin cu
ring.

c
cng thc m hnh ca s thot hi nc ni trn, nu bit E
P
, l c th tm
ra l i.
Ngo ra, c,
chng hn n c. Mt
khc, gy ot hi nc, chng hn nh bin php
hn ch chi
hi cn c vo m hnh qu trnh sinh l ca hot ng thot hi nc c ch
hot vn vn cn phi ch i nhng nghin cu
nh

Khi W < W
C
th lng thot hi n c quan h vi nng lng bc hi n
trong t,
c trong


14










iu khin s thot hi n
T
ng nc cn thit duy tr E
T
. Cn c vo lng nc ny tnh ra lng t
i p dng bin php i vi cc qu trnh khc nhau ca s thot hi n
h ph mng mng h thp D
1
s c th c ch s thot hi n
nh hng i vi E
P
h thp s th
u nng, l c th c.
K
ng thot hi nc, c mt s
ra cc thng s. Trn thc t, quyt nh vic ti phi xt n c s bc hi nc
mt t, ngha l do lng bc - thot hi quyt nh.







0,5
0
0,2
0 5 10 15 0 0,5 1,0
W, % E
P
mm/gi
0,6
a
1,0
1,0
) b)
10
6
2
0
10
0,5
1,0
0 4 8 12 16
W, %
c)
Quan h nng lng c hi n c trong t v thot hi nc
a) Quan h nc trong t v s tho hi nc (ch s l nng lng bc hi nc);
b) Quan h ca b i nc v tho i n s l lng nc trong t);
c) Quan h ca t s bc hi nc (E
P
/E
T
)
V theo hnh .2; 2- Gii gn ng)
P

E
T
/
E
Hnh 51.2. ca b
t
t h c h c (ch
(1- 45
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

TM TT
Chng ny trnh by tng i t m v cu trc v chc nng ca h sinh
thi ng rung. Vn cn a ra tho lun l cc dng nng lng dng
nhit v nc, nh sng c quyt nh quan trng n cu trc ca h thng.
Bn cnh t, qun th sinh vt cy trng cng nh dch hi u c t chc
v cu trc c trng cho tng qun x. Mi trng khng kh cng c tc
dng ng k n s pht trin ca qun th cy trng v kh nng hp th
nng lng bc x ca mt tri.
quan st v nh gi cu tr ng ca h sinh thi ng rung c v chc n
hng lot m hnh v phng trnh ton hc c xy dng v m phng
theo cc tnh cht ca chng. M hnh thot hi nc v cn bng nc c
s dng lm v d cho cc qu trnh m phng .
CU HI N TP
1. Trnh by khi nim v cn bng lng nhit ca ng rung?
2. S khc nhau gia bc x mt tri v bc x dng trong quang hp?
3. Trnh by mt cch khi qut v s phn b a l v cn bng lng nhit?
4. Trnh by phng trnh cn bng nc trn ng rung?
5. t gm nhng thnh phn g?
6. Trnh by ngun gc x ng v sinh vt gy bnh trong
t?
rng?
ng?
quan trc ca phng php kh ng lc hc
12. rnh by cc m hnh ca qun th cy trng?
n cu quang hp ca


ut hin ca cc cn tr
7. Hy trnh by cu trc sinh hc ca qun th cy trng?
8. Hy nu c im ca h thng l v h thng thn ca qun th cy t
9. Hy nu c im vt l ca tng khng kh gn mt t?
10. Hy nu c im phn b kh hu trong v ngoi qun th cy tr
11. Hy trnh by c th ca cch
v phng php cn bng lng nhit?
Hy t
13. Ti sao ngi ta li s dng m hnh khuch tn nghi
qun th?
14. Hy m t s bin i trong ngy ca quang hp qun th?
15. Hy nu mi quan h ca quang hp qun th vi chiu sng v tc gi?
70
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

16. Hy m t hiu sut s dng nng lng v h s thot hi nc?
Hy m t kh nng quang hp cc tng khc nhau trong qu 17. n th?
hp thun?
20. nh lut phn phi cht kh?
ng rung? Phng php tnh nng
trng?
28. ng ca qun th cy trng?
, thn, l ?
a nc?
i v Munekate Ken, 1972
Bn dch ca
Ni. 1998.
n l ti nguyn
v ci thin cuc sng ca ngi dn trung du min ni Vit Nam. NXB
Chnh tr Quc gia.
5. Vn Chnh, Nguyn Hi, Hong Vn Ma, Phm
p,
18. Th no l sut sinh trng tng i v sut quang
19. Vai tr ca h phng trnh sinh trng?
Hy nu
21. Hy lit k mt s loi c di trn ng rung Vit Nam?
22. Hy m t qu trnh pht sinh c di v s lng c di trn cc loi t khc
nhau nh th no ?
23. Hy m t s cnh tranh gia cy trng v c di?
24. Hy m t s cnh tranh gia cy trng v cy trng?
25. Th no l nng sut ca h sinh thi
sut sinh hc?
26. Hy trnh by tnh khu vc ca nng sut sinh hc ca cy
27. M hnh ha mi trng CO
2
trong qun th cy trng?
M hnh ha sinh tr
29. Hy m t qu trnh vn ng ca nc t t qua r
30. Th no l s thot hi nc v nng lng bc hi c
31. Hy trnh by cc qu trnh iu khin s bc hi ca nc?
TI LIU C THM
1. Ota Keizaburo, Tanaka Ichir, Udagawa Taketosh
Sinh thi hc ng rung. NXB Nng nghip, H Ni 1981 (
on Minh Khang).
2. Trn c Vin. Sinh thi hc nng nghip. NXB Gio dc. H
3. Trn c Vin (Ch bin). Thnh tu v thch thc trong qu
H Ni, 2001.
4. Masae Shiyomi and Hiroshi Koizumi. Structure and Function in
Agroecosystem design and management. CRC Press. New York. 2001.
Nguyn Mi, Trn
Thanh Nga, o Chu Thu. Gio trnh th nhng hc. NXB Nng nghi
H Ni. 2000.
71
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

6. Nguyn Tt Cnh. Nghin cu m hnh m phng ng thi m t v
chn on nhu cu ti cho ng v u tng trn t bc mu ng Anh v
ph sa sng Hng, Gia Lm. Lun n Tin s Nng nghip. HNNI, H Ni.
7. bin
8.
10.

2000.
Nguyn Mnh Chnh v Mai Thnh Phng. C di trong rung la v
php phng tr. NXB Nng nghip. 2004.
H Th Hin. Cc bin php phng tr su bnh, c di. NXB Vn ha dn
tc. 2003.
9. Phm Hong H. Cy c Vit Nam. Montreal, Php. 1993.
Nguyn Hng Sn. Mt s nghin cu v c di trn rung la cy v bin
php phng tr ng bng sng Hng. Lun vn Tin s Nng nghip.
2000.
72
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

85
Chng III
S VN NG CA H SINH THI NG RUNG
Ni dung
Trng trt bt u t vic tr c trn ng rung, thc y v c ho cc cht hu
c trong t. Nh th c ngha l ng rung lun lun gi giai on ban u ca hng
lot din bin thin nhin, ng thi, cht hu c m thm thc vt thin nhin tch lu
hng chc nm, hng trm nm ang trong qu trnh b tiu hao. Chng ny s bn v
c trng ca din bin ng rung, nhng thay i hnh thc sn xut duy tr kh
nng sn xut ca ng rung v quan h gia nng sut cy trng vi vng thch ng
sinh thi.
Cc ni dung sau y s c cp trong chng ny:
1. Din bin ca ng rung.
2. S bin i hnh thc sn xut nng nghip v ngha sinh thi ca n.
3. Trng cy thch hp vi vng t trng.



a. Nghin cu ng np lai
v thu ng 2004
b. Nghin cu ci tin
ging u tng
c. Sinh vin lm th nghim
trong nh li
nh 1.3. Mt s nghin cu v ci tin ging cy trng
ca khoa Nng hc, trng i hc Nng nghip I
(Ngun: http://www.hau1.edu.vn/khoa/nonghoc/images/khoa_hoc)
Mc tiu
Sau khi hc xong chng ny, sinh vin cn:
1. Hiu c s din bin trn ng rung v dinh dng t, thnh phn cy
trng v c di.
2. Nm c s thch ng ca cy trng vi vng t trng.
3. Nm c mt s phng php iu khin dinh dng, thnh phn cy trng
v c di trn ng rung.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

86
1. Din bin ca ng rung
1.1. S bin i m tng s ca t ng rung v s cn bng vi sinh vt
Cng vi nhng din th sinh thi ca thm thc vt t nhin, lng m tng s
trong t dn dn tng ln. Ngi ta thy rng, t tr nn mu m r rt khi din th
thm thc vt t nhin trn nhng cht phun ra t ni la, t t trng n qun th
thc vt n nh nht (climax), rng l rng thng xanh. Trong qun th rng n nh
nht c th tch lu m nhiu n 1 kg/m
2
. Do khai khn rng v ng c lm cho
trng thi cn bng gia thm thc vt thin nhin v t b ph v, din bin ca t
theo chiu ngc li, dn n phn gii v tiu hao cht hu c, trong iu kin tc
ng ca con ngi s chuyn sang trng thi cn bng mi. Hnh 1.3 cho thy s bin
i hm lng m trong t ca nhng ng c khc nhau sau khi khai khn v qua
canh tc nh nhau. Lng m tng s trong t gim xung rt nhanh t khi bt u
canh tc n nm th 10, sau c xu th chm dn v sau 40 nm th n trng thi n
nh.
t m thp qua ci to bin thnh t cn cng
c xu th bin i ging nh vy. t la nc sau
khi thot nc, lng m tng s, hiu qu phn
gii cht hu c t cn v t l amn ho cng
gim dn tng nm, qua 7 - 10 nm s gn bng cc
tr s ca t cn. Thi gian u rung nc ci to
thnh t cn, nhiu cht hu c d phn gii c
tch lu khi t t s phn gii thnh cc cht v
c m hiu qu m i vi la ht sc r rng
(Harada, 1963). T t hoang bin thnh ng
rung, hay trong qu trnh t t t bin thnh t
cn, cht hu c trong t gim i l c trng ca
din bin ng rung, nhng trong iu kin bn
phn vn khng th ni nht nh s dn n gim
ngay nng sut. Trong t t c kh nhiu cht
hu c cha mc nt, gp nhit cao s phn gii rt nhanh, v s kh oxi ca t
thng thng gy nn mc r v to thnh nhiu NH
3
- N, cy ht cht dinh dng
khng cn i v b . Sau khi ci to thnh t cn, cht hu c d phn gii gim i,
tnh trng dao ng nng sut do thi tit tht thng cng gim tng ng, v ri nng
sut n nh. Nhng s gim cht hu c trong t vt qu mt gii hn nht nh,
thng dn n gim nng sut, v cht hu c trong t l ngun cung cp cc loi cht
dinh dng v c cn thit cho cy trng sinh trng; trong iu kin thng thong kh,
chua v lng nc ph hp th hiu qu phn c bit r rt. Mt khc, thy canh
cng thu c nng sut cao, nn c nhng s nhn nhn khc nhau v ngha ca cht
hu c t. Tp on sinh vt t gm v s vi khun, si nm v ng vt nguyn sinh.
Ngun dinh dng v nng lng ca nhng vi sinh vt ny l cht hu c ca t, do
0,25
Colby
Garden City
Hays
50 40 30 20 10
0,20
0,15
0,10
0,03
0
S nm canh tc
Hnh 1.3. Bin i N trong
ng rung qua nhiu nm
G y hi ch: Hays, Colby, Graden, Cit
l a danh ca bang Kanzat
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

cht lng ca cht hu c trong t lm thay i vi sinh vt. S gim nhanh
cht hu c do din bin ng rung v vic b p li bng cch s dng qu nhiu
phn ho hc u c th ph v nhng loi cn bng no gia nhng vi sinh vt,
khin cho su, bnh ly lan trong t c xu th tng ln.
Trong t c rt nhiu tuyn trng v nm bnh lan truyn, nhng cng c nhiu loi
vi sinh vt chng li chng. Nhng loi trc tip k sinh trong nm gy bnh v dit cht
chng, bit c c: Coniothyrium minitan i vi Slerotinia, Trichoderma lignorum,
Papulospora; Penicillium vetmiculatum i vi Rhizctnia solani; Trichoderma i vi
Armillaria mellea, mt loi nm dng tuyn i vi tuyn trng... Khi trng mt loi cy
lin tc th nng sut gim r rt, mt trong nhng nguyn nhn chnh l s ph hi ca
tuyn trng v nm bnh lan truyn trong t. Th d, khi trng la cn lin tc c mt
loi tuyn trng ph hoi, qua lun canh c th trnh c s ph hi . Nhng cng c
ni c th trng lin tc mt loi cy. t ca vng quen trng lin tc tng c tng
dy, hn na phn nhiu c hm lng nc kh cao; cht hu c t, k c phn chung
bn vo c tc dng trnh hoc lm gim nh tc hi ca vic trng lin tc. Hiu qu
y l c th cung cp dinh dng v c n nh cho cy trng, ngoi ra l tc dng ca
sinh vt - pht trin nhng loi vi sinh vt v ng vt nh no c tc dng tiu dit
hoc hn ch s pht trin ca tuyn trng.
Ngoi ra, bn phn hu c cn c th phng tr c hiu qu i vi Rhizoctonia
solani, pht trin c khun Fumarium c th lm tan cht v cng trn Fusarium
oxisporum (nm bnh ho r vng), do m c tc dng c ch nm bnh. Cc vn
ni trn u l s li dng tc dng khng sinh v cnh tranh ln nhau ca vi sinh vt
ly cht hu c lm mi gii. T cho thy, ngha ca cht hu c ca t ng
rung khng ch ch l ngun dinh dng trc tip ca cy trng, ng v quan im
cn bng sinh vt, cn cn c nghin cu nhiu hn na.
1.2. Din bin ca cy trng
Nh trn ni, m tng s ca t bin thin theo thi gian - din bin ca t,
cng biu hin v mt din bin cy trng. Trong trng hp ny, din bin ca cy
trng do s la chn ca con ngi to nn. Din bin ca cy trng dn n din bin
t. Trong qu trnh bin t hoang thnh t thuc, tnh cht l ho hc ca t pht
sinh bin i, t dn n din bin cy trng. t khai hoang thi gian u mi khai
khn nhiu cht hu c v c v c ho do hiu ng t cn, d dn n t thiu xi,
nn trc ht trng la cn, khoai s v mch en l nhng cy chu c tnh trng
thiu oxi; i t thuc dn, s phn gii cht hu c gim i, mi trng cc cy cn
tng i nhiu oxi hn nh ng, i mch, c ba l v khoai ty. Nu thi gian u
mi khai khn bn nhiu phn chung v phn ln, tin hnh ci to t, th s bin i
ca cy trng s quyt nh ch yu s thay i qua cc nm v v c ho cht hu c
t. Bng 1.3 l tnh hnh bin i nng sut cy trng sau khi dng nhiu phn ln v
phn chung trn t xm ni la trung tnh. Th d, nm u sau khi khai khn t
87
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

88
thi k m phn gii ra tng i t th cy trng thch ng l khoai ty. Thi k cht
hu c ca t v c ho dn dn tng ln th cy trng thch hp l u tng. Thi k
cung cp cht dinh dng tt nht th trng ng v cy ny c tnh chng kho.
Bng 1.3. Quan h gia nng sut cy trng (kg/a) v s thun thc ca t qua cc nm
u tng Khoai ty Ng
Cy trng
Nm
Phn bn
1 2 3 4 1 2 3 4 1 2 3 4
Phn tiu chun 17,1 26,7 25,1 27,0 82,5 63,3 70,3 86,0 56,6 50,1 58,1 69,0
Phn tiu chun
+ Phn chung
23,6 31,8 30,1 27,0 96,7 81,3 65,8 84,0 61,7 61,8 65,8 69,0
Phn ln 3 ln 26,9 29,4 28,5 28,0 106,1 85,5 75,1 86,0 56,7 58,0 67,7 70,0
Phn ln 3 ln
+ Phn chung
32,2 29,0 31,7 32,0 109,3 74,5 68,8 89,0 62,2 64,0 71,6 70,0
Tnh trng cht dinh dng t thay i rt ln do lng phn bn, dn n thay
i ging cy trng tng ng. Lng phn bn, nht l phn ho hc qua tng nm,
tng ln r rt. Nh lng phn ho hc dng Nht Bn, phn m nm 1928 l 36,2
kg/ha, n nm 1975 tng ln ti 124,6 kg/ha. Hin nay vn tip tc tng ln. Tnh hnh
ny Vit Nam cng tng t. Thch ng vi lng phn cao nh vy, mt s ging
chu phn, chng v chng bnh kho c to ra. Qua cc thi k, khuynh hng
chn v bi dc ging la nc l t l thc/rm r v h s kinh t cao (hnh 2.3).
ng ch l trng lng thc tng ln qua cc nm, cn trng lng rm r th hu
nh khng thay i. Phn liu cao lm tng din tch l v cng quang hp, do
nng cao nng sut cht kh, nhng mt khc cng d lp . khc phc mu
thun sinh ra do phn liu cao, ngi ta tin hnh ci tin ging, kt qu xut hin
mt s ging c t l thc/rm r ln, chng lp v chng tt hn.







V IV III II
100
110
120
130
140
90
%
I
Khi lng ring thc
V III IV I II
90
140
%
130
120
110
100
Khi lng rm r
V III IV II I
140
%
130
120
110
100
T l thc / rm r
90
Thi k Thi k Thi k


Hnh 2.3. S bin i tnh cht ging cy trng qua cc thi k
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

89
Ngoi ra, nhng cht tit ra t r cy trng, cht phn gii ca xc hu c hoc su
bnh ly lan trong t... cng c th gy ra din bin cy trng. y s ni thm v tc
dung tng h ho hc (allelopathy). Tc dng tng h ho hc tc l nhng cht sinh
ra trong qu trnh trao i cht ca thc vt ny c nh hng ti s ny mm hay sinh
trng pht trin ca thc vt khc, quan h tng h gia thc vt vi nhau nh vy
gi l Allelopathy. l mt trong nhng nguyn nhn dn n din bin cy trng.
Th d in hnh l i mch.
Overland (1966) trng ln i
mch v Stellaria media, pht hin cht
tit ra t b r i mch c tc dng c
ch r rt i vi s sinh trng ca
Stellaria media (hnh 3.3). Tc dng
ca cht ny khc nhau i vi tng
loi cy. Nh c ch r rt i vi
Stellaria media, c ch rt nh i vi
thuc l, hon ton khng nh hng
i vi la m. Cht ly c t cy
sng v r sng c hiu qu c ch ln
hn ly t cy cht.
Theo mt s ti liu, cht ny l t
r hay thn b cht ca c mo g
(Agropyrum repens) khi nng cao,
c th c ch ny mm ca ci du v yn mch, nhng nng thp li c tc dng
kch thch (Numada, 1971).
2 6 10 14
18
0
20
40
60
80
100
Ngy
S.media + i mch
Thuc l + i mch
Thuc l
S.media
Stellaria midia
Thuc l
Stellaria midia + Cht ly c t r i mch
Thuc l + Cht ly c t r i mch
Hnh 3.3. nh hng ca cht ly c t r
i mch i vi s ny mm ca cy thuc l
v Stellaria media (Overland, 1966)
Nhng th d trn cho thy, din bin thm cy ca ng rung ngoi din bin ca
t ra, cn c tc dng sinh hc ca bn thn cy trng.
1.3 Din bin ca c di
gi ng rung trng thi ban u, hng nm cn tin hnh cy t, xi t,
lm c ngn chn din bin t nhin. nh hng ca nhng hot ng nh vy c
phn nh trn s din bin c di trn ng rung. Trn t mi khai khn, thc vt
di rng nh loi cy thn r, thn ngm, cy dy leo, hoc cc loi cy c lu nm
ca t hoang chim u th. t cng thuc dn th cc loi c hng nm cng tr thnh
s nhiu.
Trong ch lun canh rung nc - rung cn, iu kin trng trt m lng
nc trong t thay i c tnh chu k, din bin ca c di rt r. Bin rung nc
thnh t cn, ri sau mt thi gian nht nh li i thnh rung nc, cch trng trt
ny kh ph bin min Bc Vit Nam trong v ng. M, Italia v Nht Bn, vng
trng rau v nui tng i nhiu b sa cng c phng thc lun canh ny.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Bng 2.3. Din bin c di trn rung nc lun canh
Thi gian trng la Thi gian trng mu
X l
Loi
c di
1 nm
quay
li
(cy)
2 nm
quay
li
(cy)
3 nm
quay
li
(cy)
Loi c
di
1 nm
quay
li
(cy)
2 nm
quay
li
(cy)
3 nm
quay
li
(cy)
Rung nc
lin tc
a nc
a m
1.166
194
1.571
169
2.237
166
a cn
a m
83
486
239
8.580
471
7.532
Rung nc
lun canh I
a nc
a m
261
71
641
138
1.680
200
a cn
a m
15
23
49
405
177
3.864
Rung nc
lun canh II
a nc
a m
581
101
1.488
144
1.727
257
a cn
a m
36
59
260
1.335
470
7.868
Bng 2.3 cho thy, khi t cn quay li thnh rung nc, so snh vi rung nc
lin tc, s lng c di pht sinh rung nc lun canh c trong thi gian trng la
v ma ng u t v sau 1 nm th dn dn gn st vi rung nc lin tc, n nm
th ba th hu nh khng cn sai khc. Loi c di ca rung nc lun canh cng thay
i qua cc nm, t l c di a m trong thi gian trng la v t l c di a cn trong
thi gian ma ng u gim v dn dn gn vi rung nc lin tc.
Khi bin rung nc thnh t cn lun canh, cng c xu th ging nh vy.
Lun canh rung nc - t cn c tc dng c ch i vi c di, ch yu do c nh
hng rt ln n ng lan truyn ht ging c di. Nht l t rung ti bng nc
ao, h, c di mc tng i t trong thi gian di. Cn t rung ti bng nc sng,
ht ging c di chy theo nc vo rung nhiu hn. Thay i phng php tr c v
phng php trng trt cng dn n din bin c di khc nhau. Trong rung nc gn
y, mt s loi c tng ln, ngi ta cho rng do b vic sc bn, dng ph bin thuc
tr c phng tr c di mt nm v do gim trng xen.
2. S bin i hnh thc sn xut nng nghip v ngha sinh thi ca n
Ly nhng bin i v phng php ti nc trng la lm th d
Rng v ng c sau khi khai khn thnh ng rung, cht hu c ca t m thm
thc vt thin nhin tch lu b tiu hao dn qua nhiu nm trng trt, mu m ca
t v th m gim i. Nng nghip, trong qu trnh lu di nh sng to ra mt s hnh
thc sn xut va trnh c thoi ho t va gi c nng sut cao, m c pht
trin. Trong , ti ngp nc trng la v ch lun canh t cn c tc dng v
cng quan trng i vi vic gi nng sut cao cho cy trng v s ti sn xut. Nng
nghip hin i ang ly lm c s pht trin. Ti ngp nc v ch lun
canh c hiu qu cao trong vic gi mu m ca t, ng thi c th gim nh v
90
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

trnh c nhng nguy hi do c di v trng lin tc, c tc dng ln v mt n nh
nng sut cy trng.
Trong nng nghip hin i, cy trng c nng sut tng r rt trong iu kin
nhiu phn do cc cht dinh dng v c cn thit cho cy trng c cung cp y .
Nhng cng v th m dn n cy tha dinh dng, to nn nguy c lp v su
bnh ph hi.
Nguy hi ca ng rung qu mu m thng pht sinh cng vi nhng nm c
thay i tht thng v iu kin kh tng nh chiu sng, nhit . Do hnh thc
sn xut hin i phi thch ng vi bin ng kh tng v ch nghin cu khng ch
hiu lc mu m nh th no, nhm hiu r hn s cn bng ng thi gia cy
trng v mi trng trn trnh thm canh cao t nng sut cao.
Phng php trng la v tnh hai mt ca cch ti nc ngp
Ngi ta cho rng c cc phng thc trng la sau y: (1) trng la s dng nc
tri; (2) trng la ti chu k, khng lin tc; (3) trng la ti ngp nc lin tc; (4)
cui cng pht trin thnh trng la cy (m) ngp nc lin tc. Nguyn nhn bin i
ca cc phng thc trng la khc nhau l thin tai, hn hn, c di v cy mc
khng u. Phng thc sn xut trng la cc ni trn th gii hin nay c khc nhau
r rt do tnh hnh x hi v iu kin t nhin. Vng ng nh Nht Bn, Trung
Quc, Triu Tin... trng la phn ln l cy ngp nc. ng Nam , n ,
SriLanca, Italia... c cy la, cng c gieo thng ngp nc. Khc vi cc nc trn,
M, xtrylia, Xurinam u trng la c gii ho din tch ln, pht trin thnh loi hnh
gieo thng ngp nc. Coxta Rica, Panama (Trung M) li dng ma nhiu (300
mm/thng tr ln) trong thi gian lm la, p dng rng ri gieo thng khng ti. Trng
la Nht Bn 99% l cy ngp nc, mt s t vng nh Kigyoku, Okayama, Saga,
Kumamoto c gieo thng rung kh (sau ny mm 30 ngy gi trng thi rung kh, sau
mi ngp nc), cng c gieo thng ngp nc (c nc ngay khi gieo). Nh vy,
phng thc trng la, t l gieo thng v cy c s khc nhau theo vng. Ni chung, tr
vng c bit ra, u tin hnh trng la trng thi nc ngp rung, l do nh sau:
(1) Ngp nc c th thc y to xanh v vi sinh vt c nh m v tng ln d
tiu trong t. S phn gii cht hu c t b c ch, mu m t tiu hao t.
(2) Trong nc ti c cc th keo hu c v cc loi mui v c N, K, Ca, Si,
Mg... v th trng thi t ngp nc, lng cung cp cao hn trng thi t cn.
Ti nc c khi dn n hin tng qu mu m, nhng c khc nhau do tnh cht
nc ti, c tnh vt l ca t v su nc ngm. Nu t khng c tng cy
th li tiu hao mt dinh dng khi trng thi ngp nc.
(3) T nhit ca nc cao, ti nc su (15 cm hoc hn) vo ma lnh c th
bo v la trnh tc hi ca nhit thp.
(4) Ngp nc c th c ch r rt c di mc, nht l loi Panicum sp. cnh tranh
kch lit vi la, ngp nc khong 15 cm hu nh phng tr c.
91
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

(5) La nc trng ngm nc d c trng lin tc my chc nm cng khng b
hi nh cy trng cn b hi do trng lin tc. La nc trng trn cn lin 2-3 nm th
nng sut thp i r rt, do mt loi tuyn trng ph hi (tuyn trng Heterodera hi
la).
Mt khc, ngp nc thng khng li cho c nng sinh l ca r la, tuy rng cy
la nc c m thng kh vn chuyn oxi t l xung r, c th nh oxi tit ra t r
thch ng vi trng thi kh oxi, nhng cng vi nhit tng ln, s kh oxi ca t
tng mnh v tch lu cng nhiu cc loi cht b kh nh metan, axit hu c, H
2
S, cui
cng lm cho la khng thch ng c v b thi r. Kt qu lm suy gim s ht cht
dinh dng ca cy la, thm ch gy ra cc bnh sinh l nh bnh la c (Straight
head), bnh kh , bnh kh m l v bnh ng thau (Bronzing).
trng thi ngp nc, xi ho ln trn mt rung nc nh khuch tn v thm
thu m b sung vo t. Oxi ho ln trn mt rung nc, do quang hp ca cc loi
to m thay i tng ngy, trong iu kin chiu sng c th t 12 -14 mg/lt Oxi ny
thm thu xung t nh nc (tc thm thu tng ng s milimet su nc
gim mt ngy). Nhng trn 95% oxi ho ln trn mt rung nc tiu dng cho vi sinh
vt sng lp 2 cm b mt, v th oxi nh thm thu qua nc cung cp cho vng r b
hn ch rt ln (bng 3.3). T cho thy, ngp nc i vi s sinh trng pht trin
ca la nc c tnh hai mt hon ton i lp nhau.
Bng 3.3. Lng oxi ho ln (mg/l) vo trong nc trong t nh thm thu qua nc
(Tanaka, 1970)
X l
Ngy gi ly mu
V tr ly mu
Ngy
19/8
13
gi
Ngy
20/8
14
gi
Ngy
21/8
6 gi
Ngy
22/8
13 gi
20 pht
Ngy
31/8
13
gi
Ngy
9/9
13
gi
Ngy
13/9
13
gi
i chng
Nc trn t
Nc trong t
(2cm)
Nc trong t
(12cm)
9,4
0,4
0,4
9,0
0,4
0,7
3,0
0,4
0,6
9,1
0,2
0,5
6,4
0,3
0,3
14,1
0,4
0,5
13,1
0,6
0,4
Nc
thm thu
Nc trn t
Nc trong t
(2cm)
Nc trong t
(12cm)
11,4
0,3
0,3
12,2
0,6
0,6
4,4
0,5
0,6
12,2
0,5
0,7
9,0
0,4
0,3
13,3
0,3
0,2
10,3
0,3
0,3
gii quyt mu thun trong cch ti ngp nc nh vy, t trc n nay
c bin php tho nc tm thi, phi rung gia v phng thi r. Thi k phi
rung gia v tng ng vo thi k sau nhnh khi tc dng kh oxi ca t rt
mnh, trc khi sp pht sinh thi r nhiu. Lc ny, ng l thi k cy la c tnh
chng chu vi s thiu nc mnh nht trong c thi gian sinh trng ca n, qua mt
92
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

s ngy rt nc ngm trong t s c th phng trnh thi r. vng m p, do tc
dng kh oxi ca t rt mnh, nn p dng rng ri bin php phi rung gia v. i
vi t m t hoc dng qu nhiu phn xanh v qu mu m dn ti chm chn, bin
php ny cng c hiu qu. Trong khi ti ngp nc, xen vo mt thi gian phi kh
rung gia v, c th khc phc phn ln mu thun do ti ngp nc gy ra, lm cho
sn xut la pht trin ln trnh cao hn.
Hai vn mi ca cch ti ngp nc
Sn xut la nc qua qu trnh pht trin: khng ti - ti gin on - ti ngp nc
- ti ngp nc + phi rung gia v, chnh l khc phc hn hn, gim nh tc hi ca
c di, duy tr mu m ca t v trnh c tc hi do trng lin tc. Nhng ngh trng
la ti ngp nc hin nay ng trc vn lm th no iu tit hiu lc phn m v tit
kim nc. Vit Nam v nhiu nuc trng la khc, thu c nng sut la cao trn din
tch t ai c hn, s dng ngy cng nhiu phn bn, nht l phn m. Trong iu kin
lng phn cao, khi iu kin kh tng (chiu sng, nhit ...) bin i tht thng, s ht
m ca la nc thng hay trng thi qu tha, do dn n lp v . nng cao
kh nng lm du i vi nhng bin i kh tng, bin php thng p dng trc y l
dng ging chu phn chng , phng tr su bnh, bn phn nhiu ln... Nhng quan trng
nht l iu tit s ht m tng ng vi bin i iu kin kh tng. Vn th hai l li
dng hp l nc. Lng nc tiu dng cho trng la l 10.000 - 14.000 m
3
/ha, dn n
mu thun ngy cng ln vi nc dng cho cng nghip v thnh ph. Do , h thng qun
l nc hp l cn c nghin cu.
Quan h gia lng nc trong t v s sinh trng pht trin ca la
Vi iu kin nc trong t nh th no th la nc mi c th sinh trng v cho
nng sut tt nht l ti quan trng ni r s din bin h thng sinh thi la nc. V
quan h gia lng nc trong t vi s sinh trng pht trin v nng sut ca la nc
c nhiu nghin cu ch r: nng sut la nc ngy cng tng theo lng nc trong
t, t ngp nc thng c th c nng sut cao nht, la thiu nc th nng sut gim,
nht l t thi k lm ng n khi tr m gp hn th gim nng sut cng r. Lng nc
cn ca la nc l 280 -310 gam, bng hoc hi nhiu hn cc cy trng cn. V s tiu
dng nc sinh l, la nc khng nht thit c bit cn ngp nc, n ch cn nc cung
cp cho tiu hao bc thot hi nc l c. Bng 4.3 ch r s bin i nng sut ca la
nc v la cn trng trng thi t ngp nc, m t v kh. iu kin nc y l
khu ngp nc c mc nc su 5 cm, khu m t c mc nc ngm -5 cm, khng ngp
nc; khu t cn iu tit nc trong trng thi c lng cha nc ng rung bnh
thng (lng cha nc ln nht) hoc gn nh th. Bng 4.3 cho thy nng sut trong
iu kin t cn hu nh hn hn so vi iu kin ngp nc. t cn, s d tng nng
sut l do gim nh hin tng thi r, thc y phn gii cht hu c t, tng s ht
m ln mt cch tng ng. iu ni r iu kin cung cp nc v dinh dng v c
c quan h ti nng sut, ngp nc khng nht thit l iu kin khng th thiu c i
vi s sinh trng, pht trin ca la nc.
93
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

94
hi nc ca la nc tng ln theo s sinh trng, pht trin t thi
k l
t v t cn
Ging iu kin nng sut
nh khi Trng lng bng
R ca la nc phn b
nng hn so vi cc cy trng
t cn. Trn 90% s r v trng
lng r la nc iu kin
t cn c phn b trong lp
t 20cm di mt t, cho nn
phm vi ht nc v dinh dng
v c tht nh hp. T hnh 4.3
c th thy: la nc trng t
cn, lng tiu dng nc
tng t mt rt ln, d vo cui
i r n rt su, mc tiu
dng nc tng mt t (0-
20cm) vn chim 80%. La cn
th ngc li, tiu dng nc
tng t su nhiu, tnh chng
hn kho. Khi lng nc trong t tng ng vi 75-100 % lng cha nc ln
nht, quang hp ca la nc s khng bin ng v nc, nhng nu thp hn 75%
th gim thp nhanh chng. iu kin nc trong t dn ti s gim thp quang hp th
la nc cao hn u tng v lc. T cho thy, la nc l mt loi cy trng d b
gim nng sut cht kh khi s ht nc v thot hi nc trng thi khng cn bng.
Lng thot
19 - 23 / 7
0,8 0
55%
28
17
0
10
20
30
40
50
58%
35
7
0
10
20
30
40
50
16 - 20 / 8
0.8 0
Hnh 4.3. Tiu dng nc cc lp t khc nhau
ca la cn v la nc trng cn (trch dn t
"trng la nc trn t cn" ca Hasegawa)
La nc
La cn
52%
20
7
32%
28
15
13
12
m ng t din tch l ln nht n thi k tr bng, mt ngy c th n 7-8mm.
La nc trng thng xuyn trng thi t cn th gim nng sut.
Bng 4.3. Nng sut la nc iu kin t ngp nc, m
(Tanaka, 1970)
Si
(g/m
2
) (g/m
2
)
Manryo
1.755
Ngp nc
m t
t cn
1.100
956
521
437
666
Kusabue
c Ngp n
m t
t cn
1.033
1.015
1.622
470
446
465
Tachiminoru
c
1.142
Ngp n
m t
t cn
945
1.724
472
534
811
La np Norin - 1
c
1.039
Ngp n
m t
t cn
960
1.634
500
498
728
Ghi ch: Manryo e l ging la nc;
Tachiminoru v la np Norin - 1 l ging la cn.
v Kusabu
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

95
t ngp nc trn di 10
ng c th thu c nng sut
bng
i i vi c nng sinh l ca la nc, m l do
la n
ng sut l do tng s ht m.
Nhng ph t (NO - N) nu gp ma tht
th
Nhng nu ly thi k lm ng lm trung tm, cho
ngy, cn cc thi k cn li d vn trng thi t cn, c
nh thng xuyn ngp nc.
T cho thy, la nc s d thng tin hnh trng trt trng thi ngp nc
khng phi v khng ngp nc bt l
c l cy r nng, trng thi khng ngp nc h gp ma tht thng d b
thiu nc, nht l trc v sau thi k lm ng, thiu nc khi lng thot hi nc
ln nht s nh hng xu n nng sut la.
Quan h ca s vn ng nc vi m trong t
Nh trn nu, la nc khu t cn tng n
n ln m y l m dng v c trong
3
ng hoc bin ng mc nc ngm, d b ra tri v mt m, do m khng n
nh. Di y s cp n vn mc nc ngm cao thp nh hng n s vn
ng nc t cn, quan h ca s bin i chu k v lng thm thu ca n vi hm
lng m dng v c trong t.



















0
100
200
300
400
500
NH
3
-N
3

n
g

mg N/100g
NO
3
-N
NH
3
-NO
N

c

m

t

r
u
L

p

t
r

n

L

p

g
i

a

L

p

d

i

C

n
g

L

p

t
r

n

L

p

g
i

a

L

p

d

i

C

n
g

Mc n m c ngm + 7c 0 cm
L

p

t
r

n

L

p

g
i

a

L

p
C

i

n
g

L

p

t
r

n

L

p

g
i

a

L

p

d

i

C

n
g

L

p

t
r

n

L

p

g
i

a

L

p
C

i

n
g

-10 cm
-20 cm -30 cm
Hnh 5.3. Quan h c gm n
trong t (Tanaka, 1970)

Lp gi
gia m nc n v m d g v c
Ghi ch: Lp trn: 0 - 10 cm
a: 10 - 20 cm
Lp di: 20 - 30 cm
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

96
nh 5.3 ni r hm lng m dng v c (NH
3
- N v NO
3
- N) ca t tng ln
r r
o nhanh chng, nu khng
bn
ng do ma, ti v
chy
c t
cn,
hm
goi

c

y

n

H
t khi mc nc ngm h thp n 20cm cch mt t. Lm h thp mc nc
ngm, tr thnh trng thi t cn, ni chung c th thc y hot ng ca vi khun
ho kh, gim bt s mt m, dn ti tng thm m dng v c trong t. Nhng phn
ln m dng v c ca khu t cn l m dng nitrat, trc khi c la nc ht,
nu gp bin ng mc nc ngm hay ma, d b mt mt.
Ngoi ra, iu kin t cn, cht hu c ca t tiu ha
phn chung hay phn rc th lng m v c s t hn khu ngp nc. y l
mt vn cn quan tm ca vic trng trt c ti nc t cn.
Trn thc t, mc nc ngm ca rung i tr thng bin
ngm, lm cho t trng thi lc t lc kh. Hnh 6.3 cho thy nh hng lp i
lp li trng thi t cn v t t (trng thi bo ho nc) do bin i chu k mc
nc ngm, tr s trong hnh l sau 30 ngy bt u lm th nghim.
C th thy m dng v mg/100 gam t kh
rong t gim r rt do
bin ng mc nc ngm.
Loi m ny b mt mt
nhiu ch yu do mc nc
ngm bin ng lm cho qu
trnh oxi ho v kh oxi trong
t xen nhau lin tip, do
dn ti mt m. V vy,
gi c hm lng m
dng v c cao hn trong t,
phi n nh mc nc ngm,
lm cho t gi c trng
thi oxi ho hay kh oxi.
Ngc li, nh ti gin on
cho thy, khi t trng thi
oxi ho v kh oxi c tnh
cht chu k lp i lp li th
s mt m s tng ln.
trng thi t
lng m v c ca t
gim xung gn thnh ng
thng theo lng thm nc, n
dng nitrat ca tng t mt di ng xung tng su v ch nng oxi thp hn, s
lng ny khng nh. Ngi ta dng
s mt mt nh trn ra, cn lng m mt do m
15
N tin hnh th nghim, khi nng oxi l
19%, bn m dng nitrat b mt i 16% do mt m, khi nng oxi l 4,2%, m bn
b mt i 52%, iu ni r kh nng pht sinh mt m tng do oxi gim.

t

n
g

p

n
t
h

n
g

x
u

m

8


1
0

n
g

n

2

n
g

y

y

m

t
h
x
u
y

n
g

t

5

n
g

n

5

n
g
y

v

y

(A)

t

2

n
g

n

8

n
g
y

v

t

c

n

t
h
x
u
y

n
g


18
amonsunfat
canxi nitrat
16

m

d

n
g

v


Hnh 6 nh ng s hu
m t
Ghi ch: m
3
- N,
(B) ) ) (F) (E (C) (D
2
0
6
4
10
8
12
14
.3. h ca bin ng c k
c nc ngm i vi m dng v c trong
(Tanaka - 1970)
dng v c l NH
3
- N + NO
lng phn bn l 7,3 mg N/100 g t kh
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

97
thi ngp
nc
mc
nc p nc khng phi l khng th thiu i vi s
sinh
i
trong t c m -
n khu ngp nc; v khi -20 cm tr ln th khng thy sai
khc
ng ngc li vi lng ht m;
ln
v gim nng sut; loi m t va v loi m t cn th khong gia.
Hm lng m dng v c trong t trng thi khng ngp nc chu nh hng
ca lng ma nhiu t v mc nc ngm cao thp, nu so snh vi trng
th t trng thi dng tc trng thi c li tr thnh trng thi m, bt li.
Ti gin on v ti mun
Nh trn nu, hm lng m dng v c ca t c th c iu tit nh
ngm cao thp v vic ti ng
trng pht trin ca la nc. Phn ny vi mt phng php thc dng iu tit
hiu lc phn, tc l ti nc tng t lm xut hin lp i lp li chu k trng thi
ngp nc v khng ngp nc, vi cch ti gin on nh vy em so snh vi cch
ti mun khng ngp nc trong thi k nhnh, bt u ngp nc t thi k hnh
thnh ng v cch ti sm ngp nc t thi k nhnh n thi k chn, ni r
nhng c trng v hiu qu ca cc cch ti khc nhau.
Hnh 7.3 cho thy hiu qu
ca ti gin on v ti mun
vi s ht dinh dng v
nng sut, tc l mc nc ngm
v lng ma lin h vi cch
ti khc nhau thi k
nhnh (tng ng vi thi k
khng ngp nc v ca cch
ti mun), chia lm bn loi sau
y: loi m t mc nc ngm
cao, ma nhiu; loi m t va
mc nc ngm cao, ma t; loi
t cn mc nc ngm thp,
ma nhiu v loi kh cn mc
nc ngm thp, ma t.
Phn loi nh vy, m v c
20cm tr xung, nhiu h
n, nh hnh 23, khi mc nc ng
c ngha. Ngoi ra, lng thot hi nc ca thi k nhnh l t 4-6
mm/ngy. Ni mc nc ngm cao l mc nc bnh qun trong khong t -7 cm n
-18 cm; ni thp l trong khong t -25 cm n -35 cm; ma nhiu th bnh qun l 5,0-
5,3 mm/ngy, ma t th bnh qun l 3,0 mm/ngy.
Hnh 7.3 cho thy, lng ht m ca loi t m t gim, loi t cn v loi kh
cn th c tng ln. Nhng lng ht Mn c chiu h
g ht ln v silic u gim tt c cc loi, r nht l loi kh cn. V nng sut,
hnh 8.3 cho thy, khng ch c quan h vi iu kin nc m cn c quan h vi
lng m bn. loi m t, th t phn: gim nng sut, nhiu phn: khng sai khc,
rt nhiu phn: tng nng sut; loi kh cn th t phn: khng sai khc, nhiu phn: b
180
160
140
120
100
80
60
40
20
N P K Si Mn N P K Si Mn N P K Si Mn N P K Si Mn
t phn
Nhiu phn
t phn
hn
Ti mun
Nhiu p
Ti gin on
Mc nc m: Cao Cao Thp Thp
Lng m Nhiu t Nhiu t
Hnh h
ng
a:
7.3. nh ng ca ti mun v
ti gin on i vi s ht dinh dng v c
ca la nc (Tanaka, 1970)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

98
h trn ti, gia nhng
thay i ny v nng quan h biu hin bng
ph c hai sau y:
2
Da vo cng th
2
Ngha l hiu qu ng
m cy la thay ng m tt nht. D on l tr s tt
nht un l nc
cng
c cho la
t v
ln a, mt kiu kh hu thch hp
cho
xut la nc v mi trng ca n pht trin trng thi n nh hn (hnh 9.3).



1,8kg/a-N
1,2kg/a-N
0,6kg/a-N
0,3kg/a-N
Ti
mun









N nu, s ht m ca cy la nc c thay i do cch
sut trong iu kin kh tng nht nh c
ng trnh b
Y = 14,42 + 6,79 X - 0,33 X
2
(1)
Y: l khi lng go lt (kg/a)
X: l hm lng m ca cy la nc thi k lm ng (g/m ).
R = 0,901 (h s tng quan)
c ny, tm ra hm lng m tt nht l 10,35 g/m .
nh mc st gn ca hm l ca cc cch ti quyt
i do cch ti so vi hm l
ny cn c quan h vi lng chiu sng nhiu hay t; ng nhin, q
l mt trong nhng con ng quan trng t n tr s tt nht ny.
S bin i cch ti v ngha sinh thi ca n
Phn ny s i n kt lun nh v ngha sinh thi ca cc cch ti n
trng ni trn. Trng la Vit Nam v c bn chu nh hng ca nhi
g ma. Kh hu Vit Nam l kh hu nhit i gi m
la nc sinh trng pht trin. Nhng nhit v lng ma khc nhau tu theo
vng v bin ng r rt theo nm. lm du nhng nguy hi do bin ng ca iu
kin t nhin - kh hn v gi rt, nh vn thng ni, ti ngp nc c tc dng quan
trng. Khng ch nh vy, ti ngp nc cn c ch s tiu hao mu m ca t,
gim c di, trnh tc hi do trng lin tc, hon thnh v sn xut nm ny li m bo
chc chn iu kin ti sn xut nm sau. V th, trng thi ny c th lm cho sn
Ti
g n
M c ngm: Cao Cao Thp Th
L a: Nhiu t Nhiu t
Hnh 8.3. nh hng ca ti mun v ti gin on
a, 1970)
110
100
90
1,8kg/a-N
1,2kg/a-N
0,6kg/a-N
0,3kg/a-N
80
in o
70
60
c n p
ng m
i vi khi lng go lt (Tanak
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

99
Th
nhiu trng thi qu tha m, hin ti
s m o
nng cao iu kin
m qu tha, nhng do cch ti ny c tc dng
n mc
no.
Hnh 9
Trng la kt hp ti ngp
nc c nhin c th lm cho sn
xut n nh, nhng cng em li
mt s tc hi nh sinh ra thi r.
Ti t cn
Tit kim nc
i r xy ra nhiu vng m p,
t m thp giu cht hu c hoc
trong rung nc c hm lng st
thp, l mt trong nhng
nguyn nhn ch yu cn tr sn
xut la. V th, thc hin phi
rung gia v (lm thot nc
ngm) phng chn thi r c tc
dng quan trng khc phc mu
thun do ti ngp nc. Kt hp
ti ngp nc v phi rung gia
v c th lm cho nng sut la
nc tng ln ch yu da vo
cung cp nc n nh v bn
nhiu phn. Da vo bn nhiu
phn c nng sut cao ch yu
b iu kin kh tng, chiu
sng v nhit hn ch. Nht l
tnh trng chiu sng v nhit
thay i khc thng theo tng
nm, thng lm cho la nc bn
ngoi cch ti gin on thc y
khc. Ti gin on c nhin c th
kh tng, do c ch lp do ht
c ch ht m tng i ln, v vy v nguyn tc ch thch hp vi iu kin trng
trt nhiu phn. Then cht ca vic tng nng sut la nc l lm th no trong iu
kin thi tit c th, n gn vi lng ht m tt nht v loi tr nhng cht c hi
sinh ra trong t. Nhiu phn kt hp vi ti gin on c th lm cho nng sut la
nc tin ti n nh v tng bc tin st gn ti trng thi n nh ca n.
Ti mun v ti t cn c th hiu l phng hng pht trin ca ti gin
on. Thot nc c th gim bt thi r, nhng s cung cp nc v cht dinh dng
v c li tr nn khng do thiu n nh. Hiu qu ca cc cch ti ny nh th
no, ch yu nht l xem s cung cp nc v cht v c c th n nh
Ti mun
phn vo
t m, vn cha c bin php thch ng n
tnh thch ng vi s thay i
Tin hnh trng la nc trng thi t cn, nu lng nc trong t gi c
trng thi lng cha nc ng rung ln nht hoc gn nh th, th c th thu c
.3. S bin i cch ti la nc v
cc nhn t lin quan vi n
Tit kim nc
T in on
Nhiu phn dn ti tha m
i g
Ng ng gia chng p nc + phi ru
Thi r
Ti
trng lin tc
i
Cung cp dinh dng khng n nh
n
ngp nc
Tc hi rt
Tc hi
Tc hi c d
Tc hi h
Khng ti
X
X
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

nng sut bng hoc thm ch cao hn trng trt ngp nc. Nhng ti t cn kh m
thch ng c vi s thay i nhit v ma, nht l cht dinh dng chy mt khi
ma nhiu, kh bo m cung cp nc khi ma. B r la tng i nng, li dng
c t cht dinh dng v nc lp t su, nht l thi k t hnh thnh ng n
tr bng vi lng thot hi nc ln, nng sut cht kh d b gim do thiu nc. S
thay i lng nc trong t tt nhin lin quan ti s ht cht dinh dng v c, nht
l ln v silic tr ln khng n nh. Ngoi ra, mi trng sinh hc ca la nc - c
di, bnh truyn nhim trong t.... do trng thi khng ngp nc (ti t cn hoc
ti mun) lu di s tr nn phc tp hn so vi trng thi ngp nc.
Ti ngp nc s d c s dng rng ri v c th c ch r rt c di v d lm
c bng tay hoc bng my. M v Italia, nh ti nc su v lun canh rung nc
- t cn phng tr c di, c hiu qu tit kim sc lao ng lm c r rt. im ny
d k thut tr c c pht trin nhy vt nh ngy nay cng khng cho php coi thng.
Cn
thut hin nay. Do cn
pht
t s loi n nh nht thch hp vi iu kin thch ng ca n - ch yu
l iu kin kh hu (nhit , ma) v t ai. Cn s phn b v nng sut ca cy
yt nh m cn chu nh hng ca lch
s c
ni v su bnh hi lan truyn trong t, m bo iu kin ti sn xut ca la
nc, tc hi do trng lin tc l mt vn . La nc trng lin tc lu di hu nh
khng dn ti tc hi nh cy trng cn. Ngi ta thy rng, iu kin ti t cn,
hi do trng lin tc ca la cn tuyt nhin khng nh i. Trong iu kin ti mun
cho ngp nc t thi k hnh thnh ng, th khng thy hi do trng lin tc. Cn khi
gieo thng rung kh (thi k u nhnh trng thi t cn) thy c tc hi ca
mt loi tuyn trng. Tc hi trng lin tc khc nhau do cc iu kin nh loi cy
trng, s nm trng lin tc, iu kin kh tng, t ai... quyt nh; khi tin hnh lu
di ti mun trng thi t cn th d c kh nng b hi.
Tm li, t ti gin on n ti mun v pht trin cho n ti t cn, tt
nhin lin quan ti nhiu vn gp phi trong qu trnh khng ti nc n ti
ngp nc nh khng n nh cung cp nc v cht dinh dng, tc hi ca c di...
Nhng vn ny cn ang kh gii quyt vi trnh k
trin thnh mt lot phng thc ti ph hp, t ti gin on n ti mun v
ti t cn.
3. Trng cy thch hp vi vng t trng
Thm cy thin thin tri qua din th t nhin, a s tr thnh qun x n nh
(climax) vi m
trng khng hon ton do iu kin t nhin qu
i tin phng php to ging, phng php trng trt v nh hng ca hot ng
x hi. C th ly vic trng la Vit Nam lm th d ni r s ci bin cy trng
ng vng t theo s pht trin ca k thut. Ng ca nc M cng l mt th d v
trng cy ng vng t trng. Mc ny s bn v nhng quan h gia nng sut cy
trng v vng sinh thi thch hp.
100
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Bin i theo vng v nng sut la ca Vit Nam
Bng 5.3. Nng sut la h thu v ng xun ti cc vng khc nhau t nm 1995-2003
Nng sut la h thu (tn/ha)
Khu vc
1995 2000 2001 2002 2003
ng bng sng Hng 4,17 5,09 4,89 5,30 4,83
ng Bc 2,80 4,03 3,62 3,77 3,96
Ty Bc 2,17 2,45 2,69 2,76 2,91
Bc Trung B 2,48 2,87 3,24 3,46 3,65
Duyn hi Nam Trung B 2,54 3,05 3,22 3,19 3,45
Ty Nguyn 2,14 2,80 3,05 2,84 3,39
ng Nam B 2,48 2,69 2,98 3,18 3,25
ng bng sng Cu Long 2,89 3,12 3,39 3,41 3,67
Nng su t la n/ha) ng xun (t
ng bng sng Hng 4,71 5,97 5,79 5,99 6,13
ng Bc 2,95 4,56 4,47 4,65 4,87
Ty Bc 3,50 4,74 4,77 4,94 5,18
Bc Trung B 3,65 4,91 4,95 5,32 5,44
Duyn hi Nam Trung B 3,52 4,36 4,49 5,07 5,13
Ty Nguyn 3,85 4,98 4,96 4,28 4,72
ng Nam B 3,52 3,95 3,94 4,15 4,23
ng bng sng Cu Long 5,16 5,26 5,04 5,70 5,66
Ngun: Tng cc thng k (2004)
mng thng Tm (1945 nay su c c ta n
k nhau gia cc vng v su t k au i
k v gia cc vng sinh thi. Bng 5.3 cho thy nng sut la bnh qun ng bng
sng ong lun cao hn cc vng khc, ph hp vi iu
kin
T sau Cch ) n , nng t la a n tng l
hng ngng, nhng s khc nng t li r hc nh theo th
Hng v ng bng sng Cu L
a hnh v t ph sa rt thun li cho sn xut la hai vng ny. Mt khc, nng
sut la v ng xun c xu hng cao hn v h thu ti nhiu vng ca c nc.
101
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

102
Nng sut la nc, trn mt hnh thi c th biu th l: s bng/m
2
t l chn
khi lng nghn ht. Munekata xt n c trng hnh thi la nc thi k tr bng,
nhit khng kh v chiu sng thi k chn, ra cng thc tnh nng sut sau y:
N S
Y = . a
v
.
N + 50.000
.
S + 500
f
.
(LB,T) (1)
Trong :
N: s bng/ Y: khi lng go lt; S: lng chiu sng bnh qun ngy
30 ngy sau khi u; T: n ng kh bnh qun ng C)
trong 3 hi tr : khi lng phin l/m
2
; F (LB,T): ch
iu ng tng hp ca LB v T so vi nng sut (tnh c bng th

dng ngc li i vi nng sut. V nh hng ca
n
ng nh nhit thp
u
go g
khc nhau theo iu kin kh tng thi k ch
m
2
;
trong tr hit kh y (
0
0 ngy sau k u; LB
s h
tnh ton); a
v
: h s ging; : ch s hiu ng ca nhng nhn t cha bit
ngoi N, LB, S, T, a
v
.
Ni chung, nng sut tnh ra bng cng thc ny tng ln theo s bng/m
2
v lng
chiu sng tng ln, trn mt a im no th gn nh l nht nh. Nhng thi k
chn, khi nhit khng kh cao (27
0
C tr ln) nu lng chiu sng mt tri iu
kin 450cal/cm
2
/ngy, th li c tc
hit thi k chn, nh hnh 10.3 cho thy, ch s nng sut ca bt k ging no
cng u thnh ng cong ti thch m im nh ng vi 20 - 22
0
C, hn na, ng
cong v pha nhit thp th h thp rt
nhanh, v pha nhit cao th h thp chm
hn. Matsusima nghin cu nh hng ca
chnh lch nhit ngy m trong thi k
chn i vi t l chn ca la nc. Kt qu
chng minh: t l chn cao xut hin tnh
hnh nhit ngy 26
0
C, m 16
0
C ( nhit
bnh qun 21
0
C), nu nhit / ngy v nhit
/m qu cao hoc qu thp, t l chn u
gim thp. Ngoi ra, nhit thch hp li
chu nh hng ca iu kin chiu sng,
chiu sng mnh thm, nhit thch hp
cng cao ln tng ng.
Nh trn ni, nhit tt nht, trong
thi k chn quyt nh nng sut cui cng
ca la nc l tng i thp, do iu
kin nhit qu cao c

nhit khng kh bnh qun
trong 30 ngy sau khi tr u
Hnh 10.3. Quan h ca nhit
khng kh bnh qun trong 30 ngy
sau tr u v ch s hiu chnh ng
su au
(a
a
u qu
n
t ca cc ging la khc nh
gi c quan h vi vic tr hi l s
chiu sng thi k chn)
(Munekata, 1967)
c tc dng ngc i vi nng sut.
Hnh 11.3 cho thy quan h ca nng sut lt vi thi k tr u ca cc vn
n, ni r nng sut la Okayama,
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

l thp hn Akaisi thuc vng
cao
g sinh thi t
kin kh tng v rng sng
owa lm trung tm, l
vng
tch
103
Maehasi nm vng ng bng m p,
nhn vo thi k tr u thch hp nht,
r rng
rt hay Morioka ng Bc. iu
c ngha l nguyn nhn tng nng
sut vi mc cao ca vng gi rt
nh ng Bc l vic s dng rung
m bo v lm cho la tr sm ln
nhiu, lm cho thi k chn kt hp
c vi iu kin chiu sng mnh
ca thng 8, hn na nhit thi k
ny ng gn vi nhit thch hp
(21
0
C). T cho thy, vng thch hp
ca cy trng khng phi l khng thay
i, m thay i do ci cch phng
php trng trt.
Nng sut ng ca nc M v vn
nc M, ngi ta da vo iu
khc nhau, tng ng chia ra: vnh ai ng, vnh ai la m ng, vnh ai la m xun,
vnh ai bng v vnh ai cy nhit i. T vng rng n vng ng c cao u c
th trng ng mt cch rng ri, nht l vng ly min trung I
hch hp
t ai ca mi t
sn xut ng chnh. Ngi ta phn
hm lng m trong t ca cc vng
trng ng ca bang Mixixipi, v nghin
cu quan h gia N% trong t, nhit vi
nng sut ng (hnh 12.3), v thy hm lng
m trong t rng v t ng c h thp
hm s m theo nhit tng ln. Nng sut
ng bang Iowa vi bnh qun nm 10
0
C
(50
0
F) l cao nht, nhng nhit tng ln
na th nng sut li thp xung v gn thnh
t l vi s gim N% trong t. l mt
trong nhng nguyn nhn khin nng sut
ng thp hn vng nhit cao hn, iu
ni r l do nhit cao, cht hu c t
b tiu hao, tt nhin gim dinh dng cho
cy trng.
Mt khc, khi nhit bnh qun nm
trn 10
0
C, d N% trong t rt cao, nng sut
ng vn thp, chng t nhit tr thnh
10
20
0,1
30
40
0,2
0,3
32
0
40
0
50
0
60
0
70
n
g

s
u

(
b
u
s

)

it bnh qun nm
0
F
0
N

t

b

n
h

q
u

n

n

h
e
l
/
a
c
r
a
)

m

t
r
o
n
g

t

(
%
H

m

l

n
g
Nh
Hnh 12.3. Quan h nng s
i m ca t v nhit
(Jenny, 1990)
Ghi ch
N ca t ng c
ut ng
v
:
Nng sut ng
N ca tng mt t rng
N ca tng di t rng
Merioka
Maehasi
Okayama (t bng
ca Chuzoku
Akaisi (vng ni ca Chuzoku)
N: 30,000
LB: 200
400
500
65
700
(kg)
0
600
550
450
Thng 7 Thng 8 Thng 9
Hnh 11.3. ng sut
v c nhau
1 2030 10 20 30 10 2030 10
Thi k tr u
Quan h gia n
thi k tr kh
(Munekata, 1967)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

104
nh ai bng (vng nhit cao
pha
Khai khn rng v ng c m ra a bn c iu kin chiu sng tt hn cho
oi ngi trong thc tin sn
xut

nng
ng nh theo v
p hn, kho
nhn t hn ch. Wallace v Brassama cng cho rng v
Nam) nu t mu m cng c th tr thnh vng ng mnh. H cn c vo cc
im nng sut cao dng nhiu phn trng dy t 200 bushel/acr tr ln hu nh u
cc vng pha Nam. Do c ngi cho rng vng nhit cao, nng sut ng thp
c lin quan vi tnh hnh su bnh hi nhiu, nc thiu do ma phn b khng ng
u. T cho thy, vnh ai ng ca nc M trn mc rt ln l b mu m
ca t hn ch chu nh hng ca nhit, m. Klages (1949) cho rng, mt cy
trng no thch ng vi vng t no thng biu hin l nng sut cao ng u
v tng i t thay i qua cc nm. Nng sut ng ca cc vnh ai ng nc M cao
hn so vi cc vng khc v t bin ng, cng tc l ng ca vanh ai ng, ng nh
qun th n nh ca thm cy thin nhin climax, gi c trng thi tng i n
nh vi mi trng.
4. S cn bng nng lng ca qun th cy trng
cy trng, tc l sn xut nng nghip hnh thnh. L
nng nghip chn ra mt s ging cy trng c kh nng li dng bc x mt
tri cao hn, ng thi nh iu tit s cung cp nc v dinh dng, tm ra con
ng li dng nng lng nh sng
mt tri vi mc ngy cng cao.
S phn b nng lng nh
sng v quang hp ca thc vt
Ngun nng lng ca sinh vt,
bt ngun trc tip hay gin tip t
lng bc x mt tri. Thc vt
mu xanh lc hp th nng lng mt
tri, ng thi li dng ho nng do
CO
2
, nc v cht v c sinh ra trong
qu trnh quang hp, to thnh cc loi
cht hu c c nng lng. Nng
lng m thc vt mu xanh lc c
nh c trong h sinh thi, c
ng vt v sinh vt d dng khc
tiu dng, bin thnh nhit v to i
mt. Cng nng lng mt tri
lp trn kh quyn l khong 1,9 cal
cm
-2
pht
-1
, gi l hng s mt tri. C
. vng v cao, cao mt tri th
quyn di, cng nh sng v th gim thp. Th d lng tng bc x mt tri vo
tit xun phn c lng my u, 40
u sng n mt t khc nha
ng cch m nh sng xuyn qua kh
0
v Bc thp hn 30% so vi gn xch o.

1
4
Thng
3
Xch o
12
Hnh 13.3. Quan h ca v v lng t
x m t i (Berliand)
0
1 2 3 4 5 6 7 8 9 10 11
2
bc t r
1) Xch o; 2) 20 v Bc;
3) 40
0
v Bc; 4) 60
0
v Bc
ng
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

105
c x
mt
kin
u sng
ca
hch ng vi c t
t, i cn sng li, cng vi nh hng do con ngi
Nng lng nh sng mt ngy m thc vt tip thu khng nhng c quan h vi
cng nh sng m cn chu nh hng ca thi gian chiu sng. Lng tng b
tri ca ngy nng (hnh 13.3) gn xch o c nm gn nh c nh (khong 650
cal/cm
2
/ngy). vng v cao, c thay i r rt theo ma, lng tng bc x mt tri
vo cui xun n ma h li nhiu hn gn xch o. Tc sinh trng v thi gian
sinh trng ca cy trng vng v cao, tuy b nhit hn ch r rt, nhng chiu
sng ngy di v nng lng nh sng phong ph ca ma h b li s hn ch ca
nhit thp, c tc dng quan trng i vi tnh n nh ca sn xut ng rung.
Yu t kh hu, nht l lng
my, c nh hng rt ln ti iu
0,5 1,0 1,5 2,0 2,5
10
30
50
60
Cng chiu sng
Hnh 14 S k nha a c i
phn ng nh s, 1963)
G
si (Querous L.) nhiu loi thc vt thn g
thch (Aceracaea) (thc vt chu rm)
chiu sng. iu kin chiu
sng vng nhit i c s khc
nhau gia ma ma v ma kh, s
gi chiu sng ca ma ma t hn
40 - 50% so vi ma kh.
Bn thn thc vt cng c th
lm thay i iu kin chi
40
20
mi trng. Cng nh sng
lt qua trong qun th cy trng
gim theo din tch l tng v cy
vn cao. Qun th la nc sinh
trng tt, thn l xanh tt, mc lt
sng gim cn 20% tr xung. C
di trong qun th cy trng cnh
tranh vi cy trng, tc pht trin
v sinh trng ca chng b s thay
i iu kin chiu sng trong qun
th cy trng chi phi rt ln.
Thc vt trong qu trnh tin
ho, khng ngng bin i t

.3. hc u gi c lo v
sng quang hp (Mos
hi ch:
Ng
C vt (c ba l , ci ng, thuc l)
Cy
Cy h
c iu kin chiu sng khc nhau trn m
qua o thi t nhin mt s lo
bi dc nh hng v mt nng hc, lm cho s la chn ca loi cng nhanh. S khc
nhau gia cc loi v cng quang hp vi cng chiu sng nh hnh 14.3, cc
cy trng nh ng, ma, c vt (Dactylis glometata)... trng iu kin ng rung
c mt tri chiu sng tt c cng quang hp cao hn mt s thc vt nh cc
cy h thch (aceraceae) sng trong hon cnh rng c s cnh tranh v chiu sng.
Ngoi ra, t vng v cao chuyn sang vng nhit i v thp, t t thp chuyn
ln ni cao, t vng tng i nhiu my sang vng tng i nhiu ngy nng cng v
, loi thc vt c cng quang hp cao hn cng nhiu.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

106
t C
3
(1[1])
gc n i nh
la m
rong rung khng nhng v
chn
trn, thc
vt t
di m
la
nc
o, t h.

Th d, cc thc vt C
4
nh ng, ma min gc nhit i trng iu kin chiu
sng mnh, cng quang hp ca chng cao hn cc thc v
, luzec hoa tm, ci ng, thuc l; hn na, phm vi thch ng nhit ca quang
hp cng cao hn. Thc vt C
4
cn c c tnh sinh l l nhu cu nc sn xut 1
gam cht kh t hn so vi thc vt C
3
. Trong thc vt ni cao thng c nhng loi c
cng quang hp cao hn, c th t 100 mgCO
2
/cm
2
h.
C di ch yu trn ng rung thng l thc vt C
4
, iu kin chiu sng
mnh, cng quang hp cao. C di s d sinh tn c t
g c tnh thch ng rng hn vi nc v nhit , thn l v r pht trin nhanh,
tc v s lng ny mm ca ht ging cao, m cn c quan h cht vi vic chng
c th li dng c y iu kin chiu sng tt ca ng rung v c cng
quang hp mnh.
Mc ht v hiu sut s dng nng lng ca qun th cy trng
Nh nu
rong qu trnh tin ho lu
t s loi thch ng c
vi iu kin chiu sng c
chn ra. Cc loi thc vt ny
trong thi gian sinh trng
tng i ngn nh m to
phin l v vn cao cy ln,
ht v c nh nng lng nh
sng vi mc ln nht.
Hnh 15.3 l s cn bng
nng lng ca qun th
trng thi sinh trng
bnh thng (ch s din tch
l ln nht l 3,5). nh sng
chiu ti trong qun th, mt
phn t tng mt qun th
phn x i, tng ng vi
17 20% ca nh sng chiu v rong sut thi gian sinh trng, chnh lch rt n

(
1[1]
) Thc vt C
3
giai on u kh CO
2
to ra PGA(C
3
) ca ng cacbon t RuDP (ribulodifofat).
Tri li, thc vt C
4
sinh ra t axit oxaloaxetic (C
4
) ca ng 4 cacbon t DEF (fotfoenola piruvat).
Thc vt C
3
c h h hp nh sng nh CO
2
di iu kin nh sng mt tri; thc vt C
4
khng c h h
hp nh sng. Trong quang hp ca thc vt C
3
c h hp nh sng xem ra l v dng, hin nay ngha
sinh hc ca hot ng vn cha r rng. V c trng gii phu hnh thi, l ca thc vt C
4
c bao
b mch c dip lc, thc vt C
3
khng c m ny. V th, vic c hay khng c bao b mch c dip lc
l mt u mi quan trng c th phn on mt cy cha bit no thuc thc vt C
3
hay C
4
. Phn
nhiu thc vt gc nhit i l thc vt C
4
nhng la nc l ngoi l, thuc thc vt C
3
.
Tr bng
Thng 8 Thng 9 Thng 10
a
1 11 21 21 1 11 1 11
20
30
%
70
60
50
40
%
n 15 hu n i cn b n lng
trong q a n ta 68)
M

c

n

n
g

l

a
t
M

c

p
h

n

x

n
g

l

n
g

(
a
)

m

c

l

t

q
u
a

(
t
)

n
g

(
o
C
)

ng ng
v ctv., 19

c (Mura
y .3. S c
un th l
h H
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

107
ates v Loomis (1965), mc ht ca l vng mu xanh da
tri
ng lng (E ) v hiu
sut
E =
S
Mc ht nng lng ca qun th tng theo s sinh trng pht trin, sau khi tr bng
khong 20 ngy, t n tr s ln nht, khong 70%. Mc lt qua v mc ht i th l
c quan h nghch, khi thn l xum xu nht, mc lt qua gim nhiu, tng ng
di 20% nh sng chiu vo. S bin i v cn bng nng lng ny c quan h cht
ch vi s tng gim din tch l, tr bng v s bin i hnh dng cy tng ng.
Nhng mc ht nng lng nh sng ca l li c khc bit r rt do bc sng ca
quang ph mt tri. Theo G
(0,4 0,5 m) l 80 - 95 %, vng mu xanh lc (05 0,6 m) l 60 - 80 %, vng
mu (0,6 0,7 m) l 80 - 90% cn vng hng ngoi (0,7 1,1 m) th ch ht
5%, phn ln nng lng nh sng mt i theo phn x hoc lt qua. iu cho thy
l l c quan quang hp c c tnh ht la chn i vi nng lng ca bc sng hu
hiu (0,4 0,7 m) tin hnh cc phn ng quang ho hc.
Trong s nng lng nh sng m qun th cy trng ht th bao nhiu nng lng
c quang hp c nh? Biu th bng hiu sut chuyn i n
s dng nng lng (E) nh sau:
K.W K.W
E =
. S

K: nhit t 1 h la c l kho t trong thi gian
sinh trng hu nh khng i; W: lng tn t kh (g/m
2
); : t sut ht
nng
nng
qu
gam c t kh, n ng 3.760 cal/g, su
thay g ch
lng; S: lng chiu sng ca mt tri (cal/m
2
/ngy).
Hnh 16.3 ch r
hiu sut chuyn i
lng (E)
tng ng vi s sinh
trng pht trin ca
qun th la nc.
Trc khi tr bng c
tr s khong 4,7%,
thi k tr bng l
6,9%, l tr s ln
nht, sau gim rt
nhanh. Ni mt cch
khc, trong s nng
lng nh sng c
ht quang hp c nh
c khng qu 7%
(hiu sut s dng
nng lng l 3,7%),
phn rt ln nng
lng ht c b mt i a bc hi nhit v phng x nhit.
1
0
2
7
8
3
4
5
6
H
i

u

s
u

t

c
h
u
y

i

n

n
g

l

n
g

(
E

)
%
g/m
2
/ngy
L

n
g

t

n
g

c
h

t

k
h

W
)

40
30
20
10 Tr bng
W
E
21 1 10 21 1 11
Thng Thng Thng 10
Ngy thng
8 9
Hnh 16.3. S chuyn i hiu sut chuyn i
qun th c (Murata v ctv, 1968)
W
la n trong
nng lng
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

108
l do
/ L ho gi dn n h thp cng quang hp.
sng li tng.
ng nh sng.
D
phi
ca ng rung
mt phn b tiu hao duy tr cc
hot
ng rung, lm thc n cho ngi v gia sc, trong
c m
Hnh 16.3 cho thy, hiu sut chuyn i nng lng sau khi tr thp i rt nhanh
:
1
2/ Cng quang hp ca bng thp nhng ht nh
3/ Gc m ca l nm ngang lm gim hiu sut s dng nng l
o , mt trong nhng phng php nng cao hiu sut chuyn i nng lng l
chn ging sau khi tr bng c t th thu nhn nh sng tt v bn phn hp l
nng cao cng quang hp.
5. S chuyn ho nng lng
Trong s nng lng c nh nh quang hp, c
ng sng ca thc vt, nh sinh tng hp v vn chuyn vt cht protit, lipit,
nhiu loi ng v s sinh trng, sinh sn ca bn thn thc vt. Trong , t l
lng h hp so vi tng lng quang hp c s khc nhau do t l gia phn ng ho
v phn khng ng ho ca qun th, do iu kin nhit , iu kin phn bn... theo
nghin cu qun th cy trng, hng nm l khong 30 40%. Trong s cht hu c
m cy trng sn xut ra c ti 70 90% b ly ra khi ng rung. Lng li ng
rung nh l rng, gc, r... rt t. Trong cc cy trng, t l gia lng li so vi
ton b lng cht hu c, t ln hn c l cy u tng, khong 30%; la cn, ng,
khoai,... ch khong 10 - 20%.
Cht hu c thu hoch t
t phn phn chung v phn rc tr li ng rung, cng vi lng cht hu c
cn li trn ng rung c vi sinh vt phn gii, bin thnh nhit v to vo khng
trung.

Lng bc x
553 x 10
3
3,72 x 10
3
3

ca mt tri



,72 x 10
3
0,93 x 10
3
0,24 x 10
Bng
Hnh 17.3. Lng tch tr nng lng c nh ca la nc
Thn

L

R


3



trong cc b phn (kcal/m
2
)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Ghi ch: Hnh 17.3. cn c vo s liu ca Tri th nghim nng nghip Nagano
thc
dng nng
ln
u sut s dng nng lng
c
t
Hiu sut
s
hin theo "Chng trnh sinh hc quc t Nht Bn" (JIBP)
(2[2])
v nhm nghin
cu nng sut lc a (PP)
(3[3])
, mu ly l ging la Mantya, thi k ly mu l 27
thng 9 (la chn), tr s biu th bng kcal/m
2
. Nng lng tch tr ca cc c quan tnh
theo lng nhit to ra khi t (cal/g) lng cht kh (g/m
2
). Nhit t chy: bng =
4.040, l ti = 3.930, l kh = 3.580, thn (k c b l) = 3.400, r = 3.330.
Bng 6.3 trnh by mi quan h gia nng sut cht kh v hiu sut s
g ca cc loi cy trng khc nhau nhm phn tch quan h ca iu kin kh tng
v sn lng cy trng trong iu kin Nht Bn. Nng sut cy trng y t n
mc kh cao so vi cc nng h thng thng. Hiu sut s dng nng lng c s
khc bit tu cy trng, ng l cao nht, u tng l thp nht. Hiu sut s dng nng
lng cao nht m cc th nghim t c nh sau: La nc 1,64%, u tng
1,13%, ng 2,18%, ci ng 1,8%. Hiu sut s dng nng lng tnh theo nng sut
kinh t th ci ng l cao nht, u tng thp nht, theo ton b lng cht kh, th
tr s ny l 1/2 - 1/3 hiu sut s dng nng lng.
Bng 6.3. Quan h nng sut cht kh v hi
ca cc loi cy trng khc nhau (JIBP/PP, 1970)
Lng NAR
(4[4])
Trng lng ht
(g/m
2
din
ch l/ngy)
(trng lng c)
(tn/ha)
dng nng
lng (%)
ht kh
(tn/ha)
Hng mc
1967 1968 1967 1

Loi Nm
cy trng
1967 1968 1968 967 1968
La 15,05 14,64 11,54 15,86 6,08 6,33 1,17 1,26
u tng 7,28 9,99 7,60 8,05 2,87 2,76 0,17 0,81
Ng 18,38 16,68 16,66 13,35 9,07 7,56 1,52 1,36
Ci ng 16,37 20,01 9,67 10,28 10,17 12,27 1,16 1,18
Ghi ch: Hiu su c ut c

u trng trt thm canh vi cc cy trng th
hiu
php trng trt hin c t 5,0% tr ln s tng thm khng c bao nhiu.

t s d ng nng lng l tr s ly trong s thi gian sinh
trng. NAR l tr s ln nht.
Trong iu kin kh hu Vit Nam, n
sut s dng nng lng ca chng c th mong mun t t 2,5 n 5,0 trn t
tt (t mu m, thm canh cao) v t 1,5 n 3,5 trn t bc mu, t c thm canh
(bng 7.3). ng thi hiu sut nng lng cng khc nhau theo thi v v loi cy
trng. T cho thy, trong iu kin kh tng Vit Nam, tr s gii hn trn ca hiu
sut s dng nng lng c th t c vi tin cc ging cy trng v phng

(2[2])
JIBP (Japan International Biological Programme): Chng trnh sinh hc quc t Nht Bn.
(3[3])
PP: Nhm nghin cu nng sut lc a
(4[4])
NAR (Net Assimilation Rate): Sut ng ho thun (cn gi l hiu sut quang hp thun)
109
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Bng 7.3. Cc gi tr cc i ca h s s dng hu hiu bc x quang hp
trong iu kin ca Vit Nam
H s s dng hu hiu
bc x quang hp
Cy trng V trng

s
kinh t
Nng lng
thu c khi t
chy 1g ch
H
t mu
m, thm
t
bc mu, t
thm canh cao canh
t kh
ng xun 3,00 1,50 0,55
La
H 0,50 2,85 1,50
3.760
ng 4,50 2,50 0,30
Xun 4,50 1,50 0,30 Ng

3.660
H thu 2,50 1,50 0,25
ng xun 2,50 1,50 0,34
u tng
g
4.100
Thu n 2,50 1,50 0,34
ng xun 2,50 1,50 0,57
Lc
H thu 2,50 1,50 0,40
3.000
Khoai ty ng xun 5,00 3,50 0,71 1.000
Ngun: Tr nh v cng tc vin, 1997
ca cc an cy , k c l t kh
ca ng rung c nh c chuyn
ho
Chc nng ca h sinh thi gm ch yu cc giai on sau:
ng nh sng, sn xut cht hu c;
u c;
v dng
thch c v qu trnh
n c H
C th c n c vo trng l ng kh c qu trng ng ch
cy trng suy ra nng lng m cy trng
theo hng v mc nh th no. Nhng chnh xc hn vn phi biu th bng
nng lng. Th d, tnh ton nng lng ca cc c quan la nc (nh hnh 17.3), c
th thy trong s nng lng c c nh, 53% phn bng, 44% thn l, 2% phn
r. Phn rt ln bng v thn l lm ngun nng lng ca ngi v gia sc, b ly ra
khi ng rung, do nu khng c phn chung, phn rc tr cht hu c li ng
rung, th ngun nng lng cho vi sinh vt t rung, ni chung, ch c di 10% s
nng lng m cy trng c nh c. Do gim bt ngun nng lng cho vi sinh vt
t, dn n lm xu dn c tnh l - ho ca t lin quan ti hot ng ca vi sinh vt
t, v nh hng ti sn xut cy trng.
6. Tun hon vt cht ca ng rung
(1) thc vt mu xanh lc ht nng l
(2) ng vt v thc vt d dng tiu dng v d ho cht h
(3) di tc dng ca vi sinh vt, cht hu c b phn gii thnh cht v c
ng. Trong , qu trnh sn xut cht hu c ca thc vt xanh l
110
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

phn
a, Mg...
Tron
t s dng suy cho cng l bt
ngu t trong khng kh. Nhng rt nhiu thc vt khng th trc
tip
i li tr axit amin to ra cho k
ch.
gii ca vi sinh vt l khng th thiu c i vi s duy tr chc nng ca h
sinh thi, hn na hai qu trnh phi gi c quan h cn bng ng thi.
Gia sinh vt v mi trng c n vi chc loi trao i cht, l mt s nguyn
t cn thit cho thc vt nh C, H, O, P, N, P, K, S, Zn, Cu, Mn, Fe, B, Mo, C
g cc nguyn t ny, thiu bt k mt nguyn t no, i sng s c th b ngng
li. Ngun C l CO
2
; ngun O O
2
, CO
2
v H
2
O; H H
2
O; N ly t cc hp cht m
ho trong t nh mui NO
3
v mui NH
4
, suy cho cng l t N
2
trong khng kh, cc
nguyn t khc u ly t cht khong trong t, tc cung cp nhng vt cht cho
thc vt chu nh hng ca lng cung cp v tnh hu hiu. Hm lng CO
2
trong
khng kh ch l 0,03% nhng do h hp ca sinh vt, hot ng ca ni la, to ra t
cc i dng, t chy than v du... m c cung cp khng ngng, do ni chung
c th cn bng c gia tiu th v cung cp cho hot ng quang hp ca thc vt.
O
2
trong khng kh ht sc phong ph, trong mi trng lc a, tr loi t km
thng kh ra, O
2
hu nh khng phi l mt nhn t hn ch. N v H
2
O mc d c
phong ph, nhng lng hu hiu trong t c th cung cp cho thc vt trc tip ht
nu khng qu t th li qu tha, d dn n gy tc hi, c nh hng rt ln i vi
chc nng ca thc vt. Lng cung cp cc nguyn t nh P, K, Zn... c s khc bit
r rt tu thuc iu kin t. mt vng nht nh, do thiu cc thnh phn dinh
dng ny m nh hng rt ln n nng sut cy trng. Nhng so snh vi N v H
2
O
th thng lng cung cp ca chng tng i n nh. Do , di y ch yu ly N
v H
2
O ni r s tun hon vt cht ca ng rung.
Tun hon ca m
C nh m t trong khng kh: m m thc v
n t m dng phn
li dng m dng phn t, ch ht m ca hp cht m ho nh mui nitrat v
mui amon. C nhng loi thc vt c th c nh m t trong khng kh, nh vi
khun nt sn cng sinh vi cy b u, to xanh, loi vi khun quang hp v vi sinh
vt d dng Azotobacter, Clostridium... Nhng sinh vt c nh c m ny c tc
dng quan trng i vi s cn bng m ca t nhin.
Khun nt sn cng sinh vi cy b u tip thu hp cht gluxit t k ch, lm
ngun nng lng cn thit kh m dng phn t, r
Lng m c nh c ca khun nt sn c s khc nhau do chng loi k ch
v dng khun. Nh hnh 18.3 cho thy, trong s nhng cy b u, luzec hoa tm, c
ba l trng c lng m c nh c tng i cao, c th vt 200 kg/ha, lc v u
tng cng c khong 50 kg/ha. Lng m c nh c ca luzec hoa tm c th
t n bng 2-3 ln lng ht ca thc vt khng phi b u ly t trong mt nm,
iu ni r vic a cy b u vo h thng lun canh c hiu qu gi mu m
ca t nh th no. Trong nhng vi sinh vt khng cng sinh, c cc vi khun d dng
c th c nh m nh Azotobacter, Clostridium, Aerobacter; cc vi khun quang hp
c nh m l Rhodospirillum, Chromotium, Chlorobium, Rhodomicrobium; to xanh
111
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

112
cao
n ter v
khong 7
ng kh l do kh thi ca cc nh my v amon bc hi t mt t. Axit
nitri
ng vt v thc vt b mt s loi nm v vi khun phn gii, qua hp
cht
c nh m l Nostoc, Calothrix. Trong s cc vi sinh vt ny, lng m c nh ca
Azotobacter, Clostridium v to xanh tng i cao. To xanh thch ng c iu kin
mi trng kh rng, nht l d pht trin trong nc. To xanh c nh m c th nng
cao hm lng m ca rung nc, c nu r t nm 1943. Sau Chin tranh th
gii ln th hai, ngi ta pht hin nhiu loi to xanh vng nhit i, c nng lc c
nh m tng i cao, tin hnh th nghim cy ging nhiu ln trn ng rung.
Kt qu chng mnh rng, khi to
xanh sinh trng pht trin tt, tng
lng ht m ca la nc, nng
sut thc tng 5-15%. nhit thch
hp cho sinh trng ca to xanh l 30
- 35
0
C v cn chiu sng y cho
quang hp. Rung nc vng nhit
i ng Nam , n ... c iu
kin thch hp cho to xanh sinh trng
pht trin. Do , to xanh c tc dng
quan trng i vi s cn bng m
ca rung nc. Vi sinh vt khng
cng sinh y nh Azotobacter,
Clostridium u li dng nng lng t
do khi cht hu c oxi ho, va kh
m, va to ra hp cht cn cho cc
khun th. V th, lng m c nh
c ca cc vi khun ny d b nh
hng ca iu kin cht hu c do r
tit ra v nhng m rng ra, p
Clostridium. Theo c tnh, nng lc c
kg/ha. Nh vy, lng c nh m ca cc vi sinh vt khng cng sinh ny thp hn
khun nt sn.
Ngoi m c nh do sinh vt ra, cn c axit nitric, amon... do ma cung cp. C
amon trong kh
g cho s sinh trng ca Azotobac
nh m ca cc vi khun ny l
c c t hin tng phng in trong khng kh v kh thi ca cc nh my. Theo
nhiu ti liu th s cung cp m nc ma dao ng t 1 - 62kg/ha/nm, phn nhiu
l di 10kg/ha.
Phn gii cht hu c v qu trnh chuyn bin: Cc hp cht m trong xc v
cht bi tit ca
amn v axit amin bin thnh mui.
Protein R - NH
2
+ CO
2
+ cc cht khc
R - NH
2
+ HOH NH
3
+ R - OH

(
k
g
/
h
a
)

L

n
g

c

c

n
h
200
150
100
0
50
C


b
a

l


t
r

n
g

C


b
a

l

u

H


L
a
n

u

t

n
g

L

c

Hnh 18.3. L g c h c
ca cy b (Erdma 95
G

250
H
o
a

t

m

hi ch: Trch dn ca Tisdale v Nelson;
Dinh dng t v phn bn
n
n, 1

9)
n
u
m
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

113
, trong t kh hn ch yu
l c c th vi khun c tc dng chnh.
Nh
Nhng vi sinh vt tham d cc qu trnh phn gii ny
c loi khun, cn trong t m t v ngp n
ng bin i sau khi to thnh amn l: (1) di tc dng ca vi khun nitrat ho,
qua oxi ho axit hu c thnh axit nitric, (2) trc tip cho thc vt hp thu; (3) cho vi
sinh vt d dng phn gii gluxit. Trong t thng kh tt, amn hu nh chuyn ton
b thnh nitrat. Nitrat ho chia lm hai giai on: qu trnh NH
3
NO
2
v qu trnh
NO
2
NO
3
. Cc qu trnh oxi ho ny c hon thnh nh hot ng ca vi khun d
dng. Di tc dng ca Nitrosomonas, amon bin thnh mui nitric di tc dng
ca Nitrobacter, mui nitric oxi ho thnh mui nitrat. Cc vi sinh vt ny li dng nng
lng t do khi oxi ho axit nit v amn tin hnh kh CO
2
, nn c th sinh sng
c ch bng cht v c. Trong qu trnh amn bin thnh nitrat, v c bn l cn oxi,
cho nn iu kin k kh, vi khun nitrat ho s khng hot ng. Tun hon m
iu kin k kh, amon c hnh thnh cng s ngng li. Trong t thng kh tt, tc
oxi ho t NO
2
sang NO
3
nhanh hn tc hnh thnh NH
3
sang NO
3
, hn na tc
hnh thnh NO
2
nhanh hn tc NH
3
, do NO
3
c tch ly trong t. Ni chung,
thc vt sinh trng trn t kh hn c bn mui nitrat th sinh trng tt hn so vi
bn mui amon. Ngc li, la nc rung ngp nc th bn mui amn tt hn.
iu ny c lin h cht ch vi cc hp cht m ch yu ca mi trng.


N
2
trong khng kh
















P
h

n
g

n

k
h

n
g

t
r
u
n
g

P
h

n

h
o

c

V
i

s
i
n
h

v

t

k
h

n
g

c

n
g

s
i
n
h

V
i

s
i
n
h

v

t

c

n
g

s
i
n
h

T
h

c

v

t

x
a
n
h

ng vt n c
ng vt n tht
Sinh vt k sinh
Xc v cht
bi tit
Cc loi nm
v vi khun
NH
3
Ho cht m
ti thm vo
M
v gi
t i do nc
S

n

p
h

m

c

a

b
i

n


V
i

k
h
u

n

N
i
t
r
o
s
o
m
o
n
a
s

N
O
2
N
i
t
r
o
b
a
c
t
e
r

Ho tan
V
i

k
h
u

n

k
h

m

NO
3
N
2
Hnh 19.3. Qu trnh ch yu tun hon m trong t nhin ca
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

114
hn ln nitrat ni trn c thc vt ht, phn cn li b mt i do ma ra tri v
tc d ng phn nitrat ho. Khi nc trong t trng thi bo ho do ma hoc ngp
nc
hn
ng
s
. u
c c
1
thc nghim, phn ra tri do ma u c tr li cho t. Nh hnh 20.3
P

, do thiu xi m sinh ra phn nitrat ho. C th nu ra mt s khun phn nitrat


ho: Pneudomonas, Micrococcus, Achromobacter, Bacillus... Chng ly O
2
t NO
2
, kh
thnh N
2
v thot ra khi t. Hnh 19.3 l qu trnh ch yu ca tun hon m trong t
nhin. Hp cht m m thc vt ht t t, mt phn nh gc r c tr li t, phn
cn li c ng vt v thc vt d dng tiu th, li bin thnh cc cht bi tit v
xc v tr li t. Nhng cht hu c ny di tc dng ca nhiu loi vi sinh vt, nm,
b kh thnh m dng hu hiu.
Trn thc t trong tun hon ny,
m b mt i kh nhiu. Phn ln p
i v gia sc theo nc thi chy ra
bin; do ti, do ma m s ln m
trong t b ra tri, cui cng chy vo
bin. S m ny, mt phn c tr li
bng phn bn c ngun gc t sn
phm ca bin, nhng phn rt ln l
dng m thc vt lc a hu nh khng
th s dng c na, ngha l b loi ra
ngoi h thng tun hon. m c nh
bng con ng sinh hc nh khun nt
sn ca cy h u v vi sinh vt khng
cng sinh, m c nh, do phng in
trong khng kh v m ca phn ho
hc c s dng b p nhng mt
mt m nh vy, nh m gi c
cn bng m ca mi trng lc a.
cn bng m trong t: S cn bng
Di tc dng ca vi sinh vt, cht h
Quan h ca vic dng phn ho hc v
m ca ng rung b nhiu nhn t hn ch
a t bin thnh cht v c hoc cht v c bin thnh cht hu c, s ht m ca
thc vt v s kh xc hu c, s mt m do phn nitrat ho v bc hi v s c nh
m ca khun nt sn hay vi khun, mt do ra tri v c do bn phn, mt do nc
v gi cun v c do ti..., quan h gia cc nhn t ny v cng phc tp. Ngi ta
dng
15
N phn tch v bin i s lng gia chng. Jansson (1963) quan st
trong khong thi gian 6 nm v nh hng ca vic bn phn m i vi cn bng
m ca t. Trong thc nghim ny, u H Lan c bn NaNO
3
, (NH
4
)
2
SO
4
nh
du bng
5
N, trng 6 nm nghin cu ng thi ca
15
N. kt qu o Na
15
NO
3
nh
hnh 21.3.
Nm u bn 500 mg
15
N vo 5,5, kg t ct, sau dng phn thng trng
trt. Trong
m c cy trng ht
m c t gi
m mt
500
15
Nmg
300
400
0
195
100
200
6 1957 1958 1959 1960 1961
Nm
nh 20.3. Quan h v bn phn ho hc
Ghi c
H
v cn bng m ca t (Jansson, 1963)
h: Trch t Soil - nship ca Black Plant Relatio
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

115
cho
15
N.
t ca la nc
v
10 nm c bin i
lng thc, trong 5 nm u th nghim lng
3, c hai u ly gii hn lng bn phn no
(
g
thy, thot u
15
N bn vo c 57% c cy trng ht, 17% to mt vo khng
kh, cn li 26% li trong t, m phn ln l m hu c ho. Nm th hai v sau,
15
N cn li c cy trng ht vi t l rt thp, ch bng 2,6 - 3,9% hm lng trong
t lc bt u trng. Vi t l nh vy th phi 20 nm mi thu v c mt na N
nh du. iu ni r mt khi m hu c ho th vic v c ho s cc k chm.
Nishikaki t nm 1955 tin
hnh th nghim truy tm vt tch bn
Th nghim chng minh, 1,4%
ca (
15
NH
4
)
2
SO
4
bn nm u c
la nc nm sau ht, nu nm no
cng bn (
15
NH
4
)
2
SO
4
th t l
cha trong la nc cng tng hng
nm. Ngha l, rung nc do c
bn lin tc hng nm bng phn ho
hc loi amn sunfat nn dn dn
hnh thnh mu m m hu hiu.
Nhng s hnh thnh mu m m
ca t quyt nh nhng quan h
phc tp v hot ng vi sinh vt vi
phn thu hoch v li ca cy
trng, v th m khng th ni, vic
bn phn ho hc hiu lc nhanh lun
lun c th dn n tng tc mu
m m ca t.
Joka t 1962 tin hnh nhiu
th nghim v nng su
ng thi m ca t khi ch dng
phn bn ho hc vi iu kin bn
phn nh nhau trong thi gian khong
tng nm nh hnh 21.3, tnh theo trng
bn phn thch hp nht l cng thc N
. u tin nng sut la n
5
. Nhng sau li xung thp dn tng nm, tr
s cao nht v trng lng thc trong 3 nm cui cng l cng thc N
2
. Mt khc, nu
tnh trng lng rm r, th bt k nm no cng tng theo lng m tng. Ni mt
cch khc la nc bn nhiu phn nng sut rm, r hng nm tuy cao, nhng dn dn
tr thnh dng thoi ho kt ht t hn.
lm r nguyn nhn, hy quan st cacbon tng s (T - C) v m tng s ca
t sau 10 nm th nghim, nh hnh 22.
cng thc N
3
), t tng chuyn sang gim. iu ni r nu nhiu nm lin bn
lp li liu phn cao hiu lc nhanh mc nht nh tr ln, tt s dn n tiu hao
mu m ca m.
T
r

n
g

l

n

t
h

c

1 2 3 4 5 6 7 8 9 10
Nm th
T
r

n
g

l

n
g

c
h

u

40
50
60
70
80
90
100
1
1
`N
0
N
1
N
3
N
N
6
10
20
4
N
2
50
60
70
80
90
100
110
N
0
1
N
N
2
N
3
N
4
N
5
Hnh 21.3. Bin i sut la nc do
c bng phn ho hc (Joka, 1962) bn lin t
nng
Ghi ch: No - khng bn phn, 0kg/ha,
N
1
- 45kg/ha; N
2
- 60kg/ha;
a N
4
- 90kg/ha; N5 - 105kg/h
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

116
t b thoi ho,
m
gi
hu ng
tng thm
gim xung. Nh bit, mu m m ca t
ch t
t tri,
tun
ho hc: bi tnh ho ta
h c t.
T - C (%)
T - N (%)
La nc bn phn liu cao lin
tc nhiu nm lm
u m tim tng b tiu hao do
hiu lc ca phn khng tip tc duy
tr c vo thi k cui sinh trng
ca la. Rung nc thng xuyn
s dng nc bn ca thnh ph c
cha nhiu m dng amon v cht
hu c, cng c hin tng tng t
nh s tiu hao mu m m ca
t do dng qu nhiu phn ho hc.
Hnh 23.3 cho thy, sau khi ti
nc bn, lng m tng s ca t
m i. 30
0
C, ngp nc 4 tun
l lng m c phn gii thm r
rt chng t m qu tha d gy hi.
Ni khc i, m dng amn v cht
g tng thm cht mn cho t, ch
Nh trn ni, mu m m ca t tng ln do bn phn ho hc nhng khi
bn vt qu mc nht nh th
2,0
T - C (tng I)
T - N (tng I)
T - C (tng II)
T - N (tng II)
N
0
1,0
2,0
12
0
N
1 N
3
N
2
N
4
N
5
c trong nc bn hu nh chng c tc d
cht hu c d phn gii.
rong tnh hnh gii phng n nh mi c nh hng r rt n nng sut cy trng.
My nm gn y, lng phn m bn cho rung nc Vit Nam vt qu 80-
100 kg/ha/v, c ni n mc 120 kg/ha/v (Agroviet, 2002). Nhng vic bn phn
chung, phn rc li gim i r rt, hu nh u c thay bng phn ho hc. Trng
trt m dng phn liu cao lch v
phn ho hc nh vy, t b suy
thoi lu di dn n nhng hu
qu kh lng trc c.
Tun hon nc: Nc di
tc dng ca nng lng m
hon trn qu t qua cc qu
trnh bc hi, ngng ng, lu
ng. Sinh vt sng trong vng
tun hon v hn ca nc v
chim gi mt phn trong lu
chuyn ca nc. Cc c tnh l,
n, t nhit cao... ca nc c quan n i nng lng tnh di ng,
ht ch vi ton b hot ng sng ca sinh v
0
0,2
0,4
(%) T-N
1953 1962
(nc trong) (nc bn)
0
20
80
ppm
NH
1953 1962
(n c trong) (nc
4
-N
t 30
0
C
bn)
40
60
Hnh 23.3. Hm l t
k
(Tr 7)

t
r
u
n
g

b

n
h

Hnh 22.3. Bin i mu m ca t
lin tc dng phn ho hc
Gh
do
i ch: Tng I: 0 - 12cm; tng II: 12 - 17cm
S
a
i

l

c
h

t
r

n


c
h
u

n

T
r

ng m trong
trc v sau hi ti nc bn
i th nghim Nng nghip Tokyo, 196
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Ma v lng nc trong t: Tng lng nc trn qu t c tnh khong
165.000 x 10
9
acr - fut
(5[5])
, trong nc ngt chim khong 7%. Phn ln nc ngt
l n
m
c ngm v bng tuyt hai cc qu t, cng tt c nc sng ngi, ao h v
nc trong t dng c trong nng nghip ch bng khong 0,08% tng lng v ch
bng khong 0,05% lng ma hng nm (bng 8.3).
Bng 8.3. c tnh lng nc ca thu quyn qu t (Ackerman, 1959)
acr - fut So vi lng ma n
Tng lng nc
Ton lng nc ngt
hng kh
t

165.000 10
9
11.000
1850
0.14
Nc ngm
Sng v h
Nc trong k
Nc trong
Thc vt v ng vt
Lng ma nm
Lng chy ra nm
10
9
8.200 10
9
118 10
9
12 10
9
6.5 10
9
0.9 10
9
89 10
9
17 10
9
124
92
1.3
0.07
0.01
1.0
0.2
Ghi ch: Acr = 0,4047 ha; fut = 0,3048
p, tun hon n ng nht l tun hon lng nc
ma m/nm, nhng khc bit rt ln gia cc
vng
cy trng th
d d
thot hi nc trng thi khng cn bng, kh khng ng kn li.

m
V mt nng nghi c quan tr
. Lng ma Vit Nam khong 1920 m
. V d lng ma cao nht ti huyn Bc Quang (H Giang): 4.802 mm, lng
ma thp nht ti Phan R - Mi Dinh (Ninh Thun): 600 mm trong nm.
i vi sn xut cy trng, s phn b ma ma quan trng hn lng ma.
vng ma nhiu, nu phn b ma ma khng khp vi ma cn nc ca
n n hn hn. Tnh khng u ca phn b ma l mt trong nhng c trng
tun hon nc, khng th trnh khi. t, vi t cch l vt ht cha nc, c tc dng
quan trng v mt lm du tnh khng u ca tun hon nc. Nc m cy trng ht
c t t (nc hu hiu) thng biu th bng tr s ly lng gi nc ln nht ca
t tr i lng nc cy trng khng ht c (h s ho). Hnh 24.3 cho thy, nc
hu hiu khc nhau ty loi t, t tht, t tht pha st gi c nhiu nc hu hiu
hn t ct hay t st. Nhng phn ng ca cy trng i vi lng nc trong t d
trong phm vi nc hu hiu cng vn c khc nhau. Hnh 25.3 cho thy, thc vt trc
khi ho, quang hp thp xung r rt, l do lng nc trong t t i, ht nc v

(5[5])
Lng nc ngm n su 3.810 m.
117
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

118
m lng n c trong t gim
(% so vi sc ch
dn n h thp quang h
khc
g, v vy mun sn xut n l
i v nhu cu nc cung cp
hn t quan trn ng ng
ng thc
trn th gii (L Vn Khoa v ctv, 1999). Nh vy vic s dng nc hiu qu trong
nng nghip s m bo an ninh lng thc ca loi ngi.




4












H
a nc ln nht)
p cng
nhau tu loi cy trng: la
57%; u tng 45%; i mch
41%; lc 33% (Inoyama v ctv.,
1961). Hnh 20 cho thy, lng nc
cn ti cng khc nhau tu loi t
canh tc.
Cn bng nc ca cc lu
vc khc nhau v lng nc dng
cho rung nc:
Ma phn b khng u v
cha chc khp vi nhu cu
nc ca cy trn nh cn c ngun nc n nh. Nht
nhiu hn, sng ngi c i nn nng nghip rung nc c
nc c lc hay khng, r rng l n
rung. Nhng nc khng ch dng cho nng nghip; thnh ph v cng nghip cng
dng khi lng nc rt ln, nn phi nm vng cn bng nc theo lu vc.
Trn th gii, din tch t c ti tng khong 36% t nm 1970 - 1990. Tng
lng nc s dng nm 1990 cho nng nghip khong 2.700 km
g nh hng n sn l
3
. Mc d din tch
c ti chim 1/6 tng din tch canh tc nhng n sn xut 1/3 sn lng l

t

c

t

t
h

t

c

t

m

t

c

t

p
h
a

t

p
h
a

c

t

m

t

t
h

t

t
h

t

c

t

m
i
n

t

t
h

t

s

t

n
h

t

t
h

t

s

t

t
h

t

s

t

n

n
g

t

s

t


Hn 4.3 ua gi t v c gi
.D ea k 1
2
1
3
0
h 2 . Q
(U.S
n h
.A. Y
a lo
rboo
i
, Agr,
s
955)
n c
20
40
60
80
100
i mch La nc
Lng nc t t (t l % so vi
lng ch c ln nht)
rong
a n
H . n h gi
K
60
80
100
40
20
nh 25 3. Qua a lng nc
trong t v quang hp
hiu l ch h s ho
20 40 60 80 100 20 40 60 80
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Vit Nam, nhn chung s lng nc v cht lng nc phc v cho sn xut
nng nghip. Nm 1995, Vit Nam c hn 5,6 triu ha ch ng c ti, 1,4 triu ha
ch ng tiu. Bng 9.3 cho thy nhu cu nc cho 7 vng kinh t sinh thi ca Vit
Nam khong 47 t m
3
vo nm 1990, 61 t m
3
nm 2000 v d kin nhu cu s dng
nc cho ngnh nng nghip 74 t m
3
(Nguyn Trng Sinh, 1998). Tuy nhin, hn hn
thnh thong vn xy ra nh vo u ma kh nm 2004 min Trung Nam B v Ty
Nguyn lm thit hi hng trm t ng do cy trng b kh ho v cht hng lot.
Bng 9.3. Nhu cu nc cho nng nghip Vit Nam (triu m
3
)
Vng 1990 2000* 2010*
Min ni v trung du Bc B 5.959,4 8.793,2 11.273,5
ng bng Bc B 7.283,8 7.750,0 9.067,5
Duyn hi 7.699,8 Bc Trung B 5.353,3 6.206,6
Duyn hi Nam B 4 Trung .583,8 6.618,2 7.610,9
Ty Nguyn 1.792,6 2.888,3 4.324,5
ng Nam B 3,622,5 4,951,6 6.719,3
ng bng sng Cu Long 2 18.398,6 23,774,8 7.340,1
Tng cng 46.976,6 60.928,7 74.035,6
Ngun: Nguyn Trng Sinh (1998) trch t Khoa v )
h thng 6-9; lng ma c n mm; ln a h: 150 ;
c trong chn nc (tr 10-20 m
nc ca rung l ng ca i, l
xun ng la l 4-7
mm/
i,
xem c khu vc rng ln th thy khng thnh vn . V lng
nc
L Vn ctv., (1999
Ghi ch: Ma m: 1920 g ma m 0-1700 mm
3
lng tiu dng n
* c tnh
h v rung ng la) l /ngy.
Lng tiu dng a bng t lng bc h ng thm
g t v lng chy mt trn mt. Lng nc bc hi ca ru
ngy, t sai khc gia cc vng. Mc d sau khi la tt c tc dng che ph, tr s
cng hu nh khng thay i. Nc chy trn b mt c th c tip tc dng l
xt nc dng cho
thm xung t th c khc bit rt ln, do tnh vt l ca t, mc nc ngm
di t, rung nc c cy lt hay khng, rung nc thng thay i t my
milimet n my chc milimet. V th lng tiu dng nc ca rung nc c s sai
khc rt ln, tu theo cch x l i vi lng nc thm. ng thi ca nc thm
xung t vn cn mt s im cha r rng, mt phn nc thm xung t c th tr
li thnh nc chy b mt v c dng lp li. Do nng cao mc s dng li nc,
lng tiu dng nc ca rung nc din tch ln s dn dn gn vi lng bc hi.
Nhng nc tn ti lc no, ni no v vi hnh thc no li tr thnh vn i vi
nng nghip. tho mn nhu cu nc cho sinh trng pht trin v vic trng trt
chm sc cy trng, ch yu nht l nhn t thi gian. Din tch rung nc ca nc ta
khong 4,24 triu hecta (din tch gieo trng 7,35 triu ha/2 v, 2003), khng tng nhiu
119
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

tch l, cu trc l, m ca kh khng vi cc iu
kin
cc
loi h l
Ch s din tch l
nh 26.3. T s thot hi nc ca cc loi cy trng khi ch s
din tch l ln nht (Uchifuji 1969)
1. Rau dip; 2. X lch; 3. Rau cn; 4. Su ho; 5. Bp ci khng cun; 6. Su l;
7. Bp ci cun; 8. Bp ci Sn ng; 9. Khoai ty; 10. Khoai s; 11. Gng; 12. t
xanh; 13. C; 14. B g; 15. Da h ; 16. La nc (1962); 17. La c (1963);
18. La nc trng cn (1962) cn (1963); 20. Ng; 21. Da
chut (c c ng;
25. C
trong hn mt thp k qua (Tng cc Thng k, 2002; Bi Th Hng v cng tc vin,
2004). Nhn v iu kin pht sinh t, t rung c th khai khn c u khai
khn ht. Mt khc, trong qu trnh khai khn lu di, h thiu nc l b hoang, kt
qu qua v s ln sai lm nh vy, cui cng mi tm c cn bng t nhin gia din
tch rung nc v ngun nc.
Thot hi nc ca cy trng v lng nc cn: Thc vt c tc dng quan trng
trong qu trnh tun hon nc. Trong nhng cht m thc vt ly mi trng th ln
nht l nc. La nc sinh trng tt, mt ngy ht lng nc khong 70 tn/ha. S
nc ht ny, c 5% dng duy tr chc nng ca cht nguyn sinh v quang hp, phn
ln s cn li bin thnh hi nc thot ra qua kh khng. Quan h gia cc nhn t bn
trong nh s thot hi nc, din
bn ngoi nh cng chiu sng, nhit, m, gi, lng nc trong t... v
cng phc tp. Phn di s cp ti quan h nh lng gia chng, y ch nu ln
s khc bit gia cc loi cy trng v s thot hi nc iu kin c ti y .





120















Hnh 26.3 cho thy quan h gia t s thot hi nc v ch s din tch l ca
y trng iu kin gn st vi sc cha nc ng rung, khi t n din tc c
H
n u n
; 19. La nc trng
ut (b mt t); 23 c leo); 22. Da ch . u tng; 24. u t
y ngy (Rubus); 26. Nho
1,6
14
1
25
26
15
12
9
2
3
21
7
11
22
4
10
13
5
17
19
20
18
8 6
24
21
1,2
0,8
0,4
2 4 6 8
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

ln nht. T s thot hi nc cn gi l lng thot hi nc tng i, l tr s tm
c do ly lng thot hi nc chia cho lng bc hi khong trng (o bng chu
o b c hi), dng phng php ny c th loi tr c nh hng ca kh tng. T
hnh 26.3 cho thy, cc cy trng c ch s din tch l t 3 tr xung, din tch l cng
ln
ho thy, tr s ny phn nhiu trong phm vi 1,0 - 1,4, khc bit gia
cc
ng Hng (bng 10.3).

th t s thot hi nc cng cao. Nhng khi ch s din tch l ln hn 3, cy trng


khc nhau s khng thay i g na, tr s bnh qun l 1,2. iu ny cho thy r trc
khi hnh thnh tng tn cy, lng thot hi nc do nhn t cy trng l din tch l
quyt nh; sau khi hnh thnh tng tn cy th li chu nh hng nhiu hn ca iu
kin kh tng.
Ngi ta cho rng: trong qun th cy trng nc v cy che u hon ton mt
t, d loi cy trng khc nhau, ch cn sut phn x (albedo) ging nhau (mu sc l
ging nhau), khng k loi hnh t nh th no, th kh nng bc thot hi nc l c
nh v do iu kin kh tng quyt nh. Nh vy, hnh 26.3 mt mc no l
nht tr vi lun im ny. Qua cc bo co v t s thot hi nc v kh nng bc
thot hi nc c
loi cy trng tng i nh (Uchifuji, 1969). Khi din tch l t n mc no
tr ln, lng thot hi nc ca cy trng c xu th c nh, l do lng chiu sng
mt tri quan h cht ch vi s thot hi nc, c hn nht nh. Din tch l nh
nhau, nhng vi tng loi cy trng c s khc nhau cht t v lng thot hi nc l
v c s khc v lng thu nhn nh sch do cu trc tng l c khc nhau, c sai khc
v sut phn x ca tng tn cy v c s sai khc v s di ng hin nhit v tim nhit
gy ra bi dng xoy khi gi lay ng v cao cy khng u.
Lng nc cn c dng lm thc o hiu sut s dng nc ca thc vt, l
lng nc cn thit sn xut 1 gam vt cht kh. Lng nc cn l t s ca lng
thot hi nc so vi trng lng cht kh, nn n cng chu nh hng ca iu kin
mi trng lin quan vi hai nhn t . Tr s lng nc cn khc nhau cc loi
thc vt khc nhau. H Hc Ng (1986) v L Th Nguyn (1984) nghin cu v tm
ra nhu cu nc ca cc loi cy trng chnh vng ng bng s
Bng 10.3. Nhu cu nc ca mt s loi cy trng vng ng bng sng Hng
Cy trng Lng nc cn (m
3
/ha) Ngun
u tng ng 2390
u tng xun 2886
Khoai ty 2594 - 2687
Lc xun 3347
Thuc l
H Hc Ng (1986)
ng 2974
Thuc l xun 2652
Bp ci 2498 - 3625
La xun 3000 - 3700
La ma 4750 - 5890
L Th Nguyn (1984)
121
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Theo L Sm (1995) v L Th Nguyn (1984), lng nc cn to ra mt
t ay i theo ma v tro V d, u tng xun yu cu 1.064
m ng h thu yu cu 876 guyn Tt Cnh, 2000). Cc nghin cu
c c Ng (1986) cho thy cn nhiu nc hn u tng ng,
L yn (1994) tm ra nh a la m i
l
3 mm/ngy). Bng 11.3 cho thy lng nc cn (mm/ngy) ca ng ng
xun
n sn phm th ng nm.
3
nc, u t m
3
(N
a H H u tng xun
Th Ngu u cu nc c a ln hn rt nhiu so v
a xun.
Nhiu cng trnh nghin cu ca Vit Nam (m Xun Hon, 1994; Nguyn Tt
Cnh, 2000) cho thy c mi quan h cht gia m t v lng nc cn bnh qun
ngy ca ng v ng xun v v xun trn t ph sa sng Hng. Theo Nguyn Tt
Cnh (1994), lng nc cn ca u tng v xun trn t tht nh (0,23 - 4,28
mm/ngy) c mc dao ng ln hn so vi mc nhu cu ca cy trn t tht trung bnh
(0,25 - 3,7
t l thun vi m trong t.
Bng 11.3. Quan h gia m t v lng nc cn bnh qun ngy
ca ng v ng xun
Lng nc cn (mm/ngy)
m tnh theo % gi m ti a
t tht nh t tht trung bnh
80 - 100 1,29 1,27
70 - 79 1,10 1,13
60 - 69 0,62 0,57
Ngun: m Xun Hon (1994)
Theo kt qu o c ca Akrron Colorado i vi 150 loi thc vt th tr s
ny thay i r 16 loi c k Kursk (Setaria) n 1131 lo i Fraseria.
Vn khc bi ng nc cn gia cc l c vt c ch ng
vn cha r nguyn nhn. Gn y, v mt quang hp ngi ta nghin cu s khc
bit gia cc lo vt v s tng trng l cht kh, dng n t trong
nhn , khin cho vn c tin trin. Nh trn nu,
cn
t ln, t 2 i c d
t v l oi th t lu, nh
i thc ng lm m
g ch tiu ca lng nc cn
g quang hp ca thc vt C
4
cao hn thc vt C
3
, nh bng 12.3, c th thy
lng nc cn ca thc vt C
4
thp hn r rt so vi thc vt C
3
, ch bng 1/2 nhng
v sao lng nc cn ca thc vt C
4
thp, th l l vn cha r rng lm. Cn c vo
thc nghim ca Shakawy v Hesketh (1965), lng thot hi nc/cm
2
/h ca ng l
thc vt C
4
khng chnh lch ln lm so vi hng dng, yn mch l thc vt C
3
.
Nhn vo kt qu ny, nguyn nhn ch yu l lng nc cn ca thc vt C
4
thp c
th l do nng lc c nh CO
2
cao, mc sinh trng tng ng cng cao, kt qu l
lm gim mt cch tng i lng nc cn sn xut cht kh.
122
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Bng 12.3. Lng nc cn ca mt s loi cy trng
ph thuc vo h thng quang hp
H thng quang hp Tn loi Lng nc cn
Thc vt C
Rau sam Portulacc
304
349
260
K chi Panicum
La min
267
Ng
Rau gin Amaranthus
4
281
i mch
La m
ch Yn m
C mo g Agropyren semicostarim
La

Bng
Mch en
C
518
557
583
678
682
368
644
487
Thc vt C
3
u
olium incarratum
ec hoa tm
Ci b trng
Ci du
Da h
Da chut
C ba l ta Trif
Luz
614
714
377
686
636
844
Ngun: Shantz v Pieme )
Mt khc, ngay trong loi cng c thc vt C
4
v cng c vt C
3
.
V.G. Alexandrop (1924) tro n cha bit thc vt C
3
v C
4
ch ng cc
loi gn nhau v mt phn ng khc nha n na,
lng nc cn l bao nhiu l ch c
dip lc. Nh Atriplex lacin i Atriplrex l thc vt C
4
c bao b mch c
dip xut 5 gam cht kh. Ngc li, A. hortense (thc
vt
isel (1927
cng mt thc
ng thi k c ra r
loi, lng nc cn ca chng c
vi m l c hay khng c bao b m
u, h
i c quan h
iatum trong lo
lc, tiu ph 1000 gam nc, sn
C
3
) khng c bao b mch c dip lc, ch sn xut 3,4 gam cht kh. Trong h
Zygophyllaceae, Tribulus terrestris c bao b mch c dip lc c lng sn xut cht
kh trn mi n v nc dng cao hn Zygophyllum fabago, Peganum harmala khng
c m ny. Alexandrop khng c lun thuyt no v tc dng v b mch c dip lc,
Maximop cho rng m ny c tc dng thc y sn phm ng ho chy nhanh v
nhiu t l ra, do c th tng nhanh qu trnh tch ly cht kh trong thn thc vt, v
suy lun, cng c quan h vi qu trnh di ng nc trong thn thc vt.
123
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Lng nc cn khc nhau theo loi, vy th gia n v tnh chu hn c quan h
g? Vn ny c nhiu ngi ch . Ngay t nm 1957 Burton thu c kt
qu thc nghim rt th v v quan h gia lng nc cn v tnh chu hn. C chn
nui (thc vt C
4
) dng phng Nam trng trong iu kin nc khc nhau, trong
mt loi c g (Cynodon dactylon) c tnh chu hn kho, lng nc cn khi lng
nc trong t khng li thp hn khi t nc. Mt loi c g (Cynodon
dactylon), mt loi Paspalum sp., c Pangola (Digitaria decumbens), thng thy tnh
chu
xt li trong mi lin h vi h thng quang hp.
h m, s
phn
hn km hn th ngc li, trong trng hp lng nc trong t khng th
lng nc cn tng thm.
Maximop cn c vo nhiu th nghim ch ra rng tnh chu hn v lng nc
khng c quan h trc tip. Parker cho rng tnh chu hn c quan h vi su v rng
ca b r, kt cu ca nguyn sinh cht, dy ca tng m sng (cutin), tnh phn ng
ca kh khng... cng khng cp quan h vi lng nc cn. Song cng vi nhng
tin trin gn y v nghin cu quang hp, lng nc cn gia cc loi c s khc
nhau, nguyn nhn ngy cng r rng. V vy, vn quan h gia lng nc cn v
tnh chu hn cn c xem
S di ng vt cht nh nc: Nc c tc dng ho tan, chuyn vn vt cht, do
vn ng ca nc m t b mt mu hay c thm mu. Phn lc a ca qu t
khng ngng b ma bo mn, a nhiu loi mui qua cc dng sng ra bin, s lng
ny hng nm n 2.491 triu tn. Ngi ta thy rng, cc dng sng ca h thng
trm tch, lng Ca chy qua kh ln; cc dng sng ca h thng macma (ni la)
th lng chy ra ca oxit silic li nhiu hn. Rung nc s d mu m hn t cn v
trng thi ngp nc c to xanh v vi khun quang hp tin hnh c n
gii cht hu c trng thi b c ch v c thin nhin b p Ca, Mg, SiO
2
v
K c kh nhiu trong nc ti. Hin tng ti nc lm giu ng rung do ng
thi ca nc rung nc, nht l ng thi ca nc ngm m ta cha bit r, hn
na cn lng ma v s phn b ma, s lng cc cht dinh dng v c ho tan
trong nc sng bin ng kh ln, nn hin nay cha th nu r v mt nh lng.
Nn nng nghip da vo nc iu kin kh hu kh, c chiu sng y , c thm
dinh dng v c cung cp theo nc ti, v th m thu c nng sut cao. Nhng
iu kin kh hu ny, ti lu di lm cho cc loi mui tch t trn mt t, thng
dn n gim nng sut cy trng. V vy, kh cc loi mui tch t trong t cn
phi c y nc. Nn nng nghip ca ng bng sng Cu Long c duy tr
nh s cung cp cc cht dinh dng v c v cc cht hu c do sng Cu Long
khng ngng mang li v tc dng ra mn ca nc l chu k.
Ngc li, vng m t nhiu, t b bo mn v tiu hao dinh dng r rt, nht
l cc vng t dc, bo mn l mt vn ln. gi nc v t, mt t cn c c
t nhin hay cc lp thm t nhin hay nhn to khc che ph, v cn bn phn hu c
hnh thnh cu trc vin bn trong nc, tin hnh trng trt theo bng v canh tc
theo ng ng mc.
124
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn


TM TT
H sinh thi ng rung c qu trnh pht trin lu di bao gm s bin i
m tng s v cc cht dinh dng khc trong t kt hp vi din bin ca
cy trng v c di. Cc phng php iu khin s pht trin ca cy trng,
hn ch s pht trin ca c di v huy ng thnh phn dinh dng trong t
c cp n trong chng ny. S lng v thnh phn loi c di c s
thay i su sc khi t c thun thc ha. c bit, phng php ti gin
on v ti mun c kh nn hm lng m v c trong t

g iu khin
thng qua s thay i mc nc ngm cao hay thp.
Khc vi s phn b ca thm thc vt t nhin, s phn b v nng sut ca
cy trng khng hon ton do iu kin t nhin quyt nh m cn chu nh
hng ca lch s ci tin phng php to ging, phng php trng trt v
nh hng ca hot ng x hi. La chn cy trng thch hp vi tng vng
sinh thi c ngha l chn cy trng ph hp vi iu kin t ai v kh hu
ca tng vng. y cng l ch trng chuyn i c cu cy trng v vt
nui ca Chnh ph Vit Nam.
CU HI N TP
1. Hy trnh by s bin i m tng s ca t ng rung v s cn bng vi
sinh vt?
2. Qu trnh din bin ca cy trng xy ra nh th no trn ng rung?
3. Hy nu din bin ca c di trn ng rung?
4. S thay i v phng ha sinh thi nh th no?
5. Hy nu mi lin h gia phng php trng la v tnh hai mt ca cch
ti ngp?
7. ca s vn ng nc vi m trong t?
i vng t trng ?
php ti nc c ng
6. Hai vn mi ca cch ti ngp nc?
Mi quan h
8. S khc bit gia ti gin on v ti mun?
9. Hy nu phng hng la chn cy thch hp v
125
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

TI LIU C THM
hi v Munekate Ken, 1972
ch ca
2. Masae Shiyomi and Hiroshi Koizumi. Structure and Function in
Agroecosystem design and management. CRC Press. New York. 2001.
3. Nguyn Tt Cnh. Nghin cu m hnh m phng ng thi m t v
chn on nhu cu ti cho ng v u tng trn t bc mu ng Anh v
ph sa sng Hng, Gia Lm. Lun n Tin s Nng nghip. HNNI, H Ni.
2000.
4. Trn c Vin. Sinh thi hc nng nghip. NXB Gio dc. H Ni. 1998.
5. Trn c Vin (Ch bin). Thnh tu v thch thc trong qun l ti nguyn
v ci thin cuc sng ca ngi dn trung du - min ni Vit Nam. NXB
Chnh tr Quc gia. H Ni. 2001.
6. Tng cc Thng k. Nin gim thng k 2003. NXB Thng k, H Ni.
2004.


1. Ota Keizaburo, Tanaka Ichir, Udagawa Taketos
Sinh thi hc ng rung. NXB Nng nghip, H Ni 1981 (Bn d
on Minh Khang).
126
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Chng IV
IU KHIN H SINH THI NG RUNG
Ni dung
Trong lch s pht trin ca loi ngi, h sinh thi ng rung khng ngng c
ci tin v iu chnh theo hng c li cho con ngi. V phng php iu khin,
ngi ta thng nhn mnh n k thut iu khin vt l, iu khin ha hc, iu
khin sinh hc (bao gm lai to ging).

nh 1.4. Nghin cu t hp ng lai 5 X TN115 ti trng HNN I (nm 2003)
Cc ni dung sau y s c cp trong chng ny:
ngha v phng php iu khin h sinh thi ng rung
iu khin qu trnh ca h sinh thi ng rung
Mc tiu
Sau khi hc xong chng ny, sinh vin cn:
Hiu c c s l lun v thc tin ca cc phng php iu khin h sinh thi
ng rung
127
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

1. ngha v phng php iu khin h sinh thi ng rung
H sinh thi ng rung trong iu kin hot ng ca con ngi khng ngng
c iu chnh theo chiu hng c li nhm cung cp nhng sn phm cn thit cho
con ngi. Trn phng din ny, n c th c gi l h sinh thi b iu khin. Tuy
nhin, con ngi ph v mi cn bng vn c ca th gii t nhin do gii hn v
khoa hc cng ngh cng nh nhn thc. Chnh v vy, nhng tc ng ca con ngi
trong sn xut nng nghip cn c nghin cu mt cch k lng ra cc bin
php khc phc hp l. V phng php iu khin, ngi ta thng nhn mnh n k
thut canh tc; ni mt cch tng i, theo quan im ci to chc nng ca h sinh
thi th cha y (Phm Ch Thnh v ctv, 1996; Trn c Vin, 1998). Nh vic ci
to bn thn cy trng, tc l pht trin k thut to ging lm nng sut tng vt,
ng nhin khng ch tho mn tnh nng cho nng sut cao m cn nng cao c tnh
chng chu su bnh v thin tai, v.v... Vn iu khin h sinh thi ng rung nu
trong chng ny ch yu l ci tin chc nng ca h sinh thi nhm nng cao nng
sut cy trng.
Phng php iu khin c th chia ra lm ba kiu: iu khin vt l, iu khin
ho hc v iu khin sinh hc.
iu khin vt l
Mi trng xung quanh cy trng c th hiu l iu kin kh tng v kh hu
trong tng khng kh gn mt t, cc iu kin ny thng c tc dng bt li i vi
s sn xut ca cy trng. c thu hoch n nh, phi khc phc cc im bt li ,
nghin cu ci thin iu kin kh tng v kh hu. V th c bit coi trng nghin
cu cu trc mi trng m quan trng l bc x mt tri (chiu sng mt tri) v h s
khuch tn. i vi bc x mt tri, tc dng ca bin php nng nghip nhiu lm mi
ch l phng nhit ln cao do chiu sng qu , hoc dng vt liu che ph
thay i sut phn x, hu nh kh gy c tc dng no tch cc hn na. Trng rng
chn gi gim tc gi, vt che chn hay b t cng c th gim tc gi gn
mt t, ngha l c tc dng i vi h s khuch tn. i vi t, nh cy ba m ci
thin tnh cht vt l ca t, ti nc c th iu khin tnh trng nc ca cy trng,
nu khng ni n tc dng no khc, th chnh l ci thin tnh trng nhit gn
mt t.
Bng 1.4 l mt kt lun nh v s ci thin iu kin nhit gn mt t. Do c
s tn ti ca cy trng, nn nhng tc dng t nhiu cng c bin i, nh hng
ca bn thn cy trng i vi nhit th khng ln, nhng s tn ti ca cy trng
lm thay i mc lt vo ca nh sng hoc lm gim tc gi, mt s nh hng nh
vy tng i r rt.
128
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

129
Bng 1.4. iu khin vt l i vi s ci thin iu kin nhit gn mt t

Tc dng i vi
bc x thun
Bin i sut
phn x
Bin i
bc x hu hiu
sng di
1. iu kin sut phn x
ca b mt t
2. Nh thay i lng nc
trong t m iu tit
sut phn x
S dng mn khi v mng
mng c tnh nng hp thu
la chn i vi sng ngn
v sng di
Tc dng i vi
tnh cht nhit
hc ca t
Thay i h s
truyn nhit v
nhit dung
1. iu tit lng nc
trong t
2. Ph t iu tit tnh
cht nhit
3. Nh cy ba m thay i
tnh cht t
Tc dng i vi
h s khuch tn
tng kh
Gim tc gi
v cng
dng gi xoy
1. Dng rng v tng t
gim tc gi
2. Dng mng vinin
cch ly vi gi
bn ngoi
3. Che chn v b t
gim tc gi
gn mt t
Tc dng i vi
bc hi nc
mt t
Thay i tc
bc hi nc
1. Che ph bng mng
vinin
2. Dng c ch bay hi
3. Ti iu tit lng
nc trong t














Ci thin
iu kin
nhit
gn mt t








iu khin ho hc
Dng cht ho hc iu khin s sinh trng pht trin ca cy trng c hai
hng: mt l tc dng ca phn bn i vi sinh l dinh dng ca cy trng; hai l tc
dng ca cc cht iu tit sinh trng m ch yu l cht kch thch vi sinh l sinh
trng pht trin ca cy trng. Ngoi ra, vic iu khin h sinh thi ng rung cn
gm c vic s dng rng ri cc cht ho hc (thuc bo v thc vt) iu khin qun
lc c di v phng tr su bnh. Vic s dng thuc bo v thc vt hin nay ang tn
ti nhiu vn do coi thng nhng hu qu bi nhng tc dng v phn tc dng
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

130
ca thuc bo v thc vt gy nn trong h sinh thi. y khng ch l vn ca h
sinh thi ng rung, nu thiu suy xt su xa n cc h sinh thi khc th vn cng
s khng c gii quyt.
iu khin ho hc s sinh trng pht trin ca cy trng phi gm nhng ni
dung sau y v cc phng thc ny c gi l iu khin ho hc i vi cy trng.
1. iu khin ny mm, ng ngh.
2. iu khin ho hc sinh trng chiu cao cy, nhnh.
3. iu khin ho hc tr bng n hoa.
4. iu ch ho hc hin tng chn v hin tng ho gi.
5. iu khin ho hc cc c tnh sinh thi: tnh chng nng, tnh chng rt.
6. iu khin ho hc cht lng sn phm.
iu khin sinh hc
Cho n nay, ni iu khin sinh hc l ngi ta ngh ngay n vic ci tin ging
cy trng, chn to ra cc ging mi. To ging c tc dng rt ln trong vic tng
nng sut la nc ca Vit Nam v th gii (Trn Cng Tn v ctv, 2002). Cng tc
nghin cu to ging gn y tin thm mt bc do to ra cc ging la lai 2
dng, 3 dng (Nguyn Th Trm, 2003).
V mt phng tr su bnh hi, vic nghin cu li dng thin ch t lu c
ch trng (Odum, 1983; Trn c Vin, 1998). Thin ch ca su hi c cn trng k
sinh, virut, vi khun, ng vt nguyn sinh..., trong , vic li dng vi sinh vt ang
c coi trng nht. T cho thy, phng hng iu khin h sinh thi ng rung,
ngoi cc phng php c ra, tin hnh iu khin bng nhng tc dng tng h
gia cy trng v mi trng ngy cng tr nn quan trng. Do , nu r tnh quy lut
nh hng v tc dng tng h gia cy trng v mi trng l ht sc cn thit.
Di y xin ly vic iu khin nhit lm th d ni r s ci thin mi
trng trng trt; i vi h sinh thi ng rung, y c th l mt v d tng i c
th. Sau s bn n nhng suy ngh v iu khin tng hp, iu khin h thng ph
v vic sn xut cy trng c k hoch.
2. iu khin qu trnh ca h sinh thi ng rung





C cu
iu khin
Tn hiu
iu khin
Qu trnh
Tn hiu
Tn hiu
+
-
Hnh 1.4. Phng thc iu khin c bn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

131
H thng iu khin thch ng
Trong sn xut cng nghip thng gp phi vn : cho sn phm ph hp vi
mt yu cu ca mt ch tiu no , phi tin hnh iu khin v o gim nh i vi
chc nng b phn ca h thng ch to sn phm , thng thng gi l vn
iu khin qu trnh, l mt lnh vc nghin cu ca k thut iu khin. Phng thc
iu khin ny l mt loi iu khin t ng, thu c thng tin t b phn ca h
thng hoc t u chuyn ra, nh m iu khin chc nng ca h thng. Hnh 1.4 l
mt th d c bn v gin n trong h thng c chc nng nh vy. l phng thc
ly thng tin chuyn ra ca h thng quy v pha chuyn vo, nh kim tra sai s tn
ti m iu tit tn hiu chuyn i. Phng thc iu khin ny c hiu sut v chnh
xc km, phm vi thch hp hp, khng thch hp dng vo h thng quy m ln. H
thng x l trong k thut h thng ni chung u c nhiu lp chuyn vo, qu trnh
bn trong phc tp v thng pht sinh tc dng qua li. Qu trnh ca h thng ny cn
c mt c tnh l bin i c thi gian theo mt s nguyn nhn pht sinh bn ngoi
hoc bn trong. Do , iu khin h thng quy m ln, phi nm vng s thay i
c tnh ca qu trnh do cc bin ng ny gy ra, ni mt cch khc l phi hiu c
c tnh ng thi ca qu trnh. H sinh thi m chng ta nghin cu hon ton c th
coi l h thng quy m ln nh vy. nm vng c tnh ng thi ca qu trnh, cn
tin hnh tnh ton v o lng nht nh thu c nhng ti liu phn on cn thit
cho vic iu khin. Nhng phn on ny c hon thnh qua my tnh in t.
dng my tnh in t vo qu trnh sn xut, t nht cn c ba iu kin sau y:
1. C nhng my o lng thch ng vi tin cy cao.
2. C h thng iu khin ch lnh tr li chnh xc v nhanh.
3. Phi c mt m hnh qu trnh tiu chun biu hin c iu kin tt nht.
iu khin qu trnh cn c m hnh (Ahuja, 2002; Phm Ch Thnh v ctv, 1996;
Tsuji v ctv, 2002). Hnh 2.4 l phng thc iu khin qu trnh khng c m hnh.
Trong phng thc ny, hiu chnh chuyn vo, s dng tn hiu chnh pht ra chu
k, ngoi vic dng my tnh th nghim ng tc ra, v c bn l ging vi hnh 1.4.
Phng php ny do khng li dng thng tin bn trong h thng, cho nn lng thng
tin ca tn hiu chnh t c cu iu khin ra rt t.






Hiu chnh Tn hiu ra
Kt qu ng tc
C cu
iu khin
Qu trnh
My tnh th
nghim ng tc
Tn hiu vo
+
Tn hiu
iu khin
Nhiu
Hnh 2.4. Phng thc iu khin qu trnh khng c m hnh
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

132
Khc vi phng thc trn, phng thc iu khin nh hnh 3.4 c th thu c
nhiu thng tin hn trong qu trnh iu khin, c th lin tc tin hnh thao tc tt
nht. im khc vi phng thc iu khin khng c m hnh l phng thc ny c
m hnh hon ton c lp v dng my tnh in t kim nghim chnh xc ca
m hnh. Do , trong h thng ny, c th va bc vo iu khin qu trnh, va
chnh l kt qu ca hnh vi mnh to ra, nh tin hnh hiu nh tu lc i vi mi
trng mnh hiu nh quyt nh tiu chun, nn cng gi l h thng iu khin
thch ng. Trong h thng quy m ln tng i phc tp, trc ht cn t m hnh
vo trong h thng, v th phi lm r cn iu khin bin s no trong m hnh, tc l
iu khin b phn no trong qu trnh, sau mi tin hnh iu khin chnh xc.
Cn nghin cu lm th no sn xut cy trng trong h sinh thi ng rung tr
thnh h thng iu khin thch ng nh vy m iu khin. Nhng hin nay vn cha
c m hnh dng thch hp vi h thng tng hp, thm ch ngay n m hnh sn xut
(sinh trng) cy trng cng khng hon thin. Di y s ni n c s ca s iu
khin qu trnh h sinh thi ng rung - iu khin quang hp ca qun th cy trng.







Tn hiu ra
Kt qu
ng tc
M hnh
My tnh
iu khin
My tnh th
nghim ng tc
Qu
t h
My tnh th
nghim m hnh
+
-
Tn hiu
iu khin
Hnh 3.4. Phng thc iu khin thch ng (Kunizawa v ctv., 1964)
iu khin quang hp ca qun th cy trng
H thng sn xut cy trng l mt h thng ph ca h sinh thi ng rung, m
quang hp qun th li l mt h thng con ca h thng sn xut cy trng. V s
quang hp ca qun th cy trng, mc d c nhiu thng tin, nhng phng php iu
khin coi n l h thng iu khin thch ng th li hu nh cha c nghin cu. M
thc m hnh ho nu ra chng trc c th dng lm m thc iu khin. T cng
thc (95) n (99) c th thy r quang hp ca qun th quyt nh :
1. Hm s nh sng - quang hp ca phin l
2. Hm s CO
2
- quang hp ca phin l
3. Hm s h hp ca phin l
4. Cu trc hnh hc ca qun th (h s tiu nh sng)
5. H s khuch tn trong v ngoi qun th
6. Bc x mt tri
7. Mt ct phn b thng ng ca nng CO
2
.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

133
Do , phng php iu khin cng phi nghin cu ln lt tng yu t.










CaO% MgO% SO
3
%
K
2
O% N% P
2
O
5
%
1
2
3
1
2
3
0 1 2 3 4 0 0,2 0,4 0,6 0,8 0 1 2
0 0,2 0,4 0,6 0,8 0 0,2 0,4 0,6 0,8 0 0,5 1,0
Hnh 4.4. Hm lng thnh phn v c v cng quang hp ca phin l la nc
Hm s quang hp ca phin l
Nhng nhn t ch yu quan trng quyt nh hm s quang hp ca phin l l:
cu trc ca l (k c cu trc ca cht dip lc), tui ca l v c th cy, thnh phn
v c (hu c) ca l, lng nc ca l v nng lc di truyn ca ging. Trong ,
thng qua iu kin canh tc c th thay i ch yu l thnh phn v c (hu c) v
lng nc ca l. Cc thnh phn v c c tc dng l nhn t hn ch i vi hm s
quang hp ca l (hnh 4.4). Nhng l kt lun rt ra iu kin trng trong nc v
trng trong ct khi thnh phn dinh dng v c ht sc thiu. Cn cc trng hp
thng thng, thnh phn dinh dng v c, nh m chng hn, ch nh hng cht t
ti hm s quang hp m thi (thc t ch nh hng ti hm lng protein). m c
tc dng quyt nh i vi s m rng din tch l, c bit quan trng i vi s iu
khin quang hp thun ca qun th thiu nc th t trc khi ho cng h
thp quang hp. Ni chung, quan h hm s quang hp v nhit c th v thnh
ng cong tt nht, nhit ng vi gn nh ng cong tng i gn vi nhit
khng kh ca thi k trng trt cy trng .
Hm s h hp ca l n
Thng thng, hot ng h hp v hot ng quang hp l song song, c ngha l,
l c hot ng h hp mnh th hot ng quang hp ca n cng mnh. V th hot
ng h hp c quan h cht ch vi tui l, l cng non th cng mnh. Ngoi ra hm
lng m cng c quan h cht ch vi hot ng h hp. Do cung cp m, hot ng
h hp v s tng hp protein lm nhn qu cho nhau v u c thc y. Nh
bit, nhit thp c th h thp hot ng h hp, nhng nhit tt nht ca h hp
li khc nhau tu loi cy trng. C cao ln 10
0
C so vi nhit tt nht, th thc y
h hp ln gp 2 - 3 ln, ging vi tr s ca nhng phn ng ho hc ni chung.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

134
Cu trc hnh hc ca qun th
Chng trc ni k vn ny, ch nhn mnh l mt qun th khc nhau
th phng thc nh hng khc nhau r rt. H s khuch tn quyt nh tc gi
v cng dng xoy. Ni chung, tc gi cng ln th h s khuch tn cng cng
ln, tc trao i (tch phn ca h s khuch tn) gn nh t l vi tc gi.
Mt ct thng ng ca nng CO
2
F
t
= 40
k = 0.5
Q
N
= 10ly/min
K
N
= 500cm
2
/g
45 50 55 60 65
C.10
2
g CO
2
/cm
3
Nng CO
2
t t thi ra c
nh hng i vi mt ct thng ng
ca n (hnh 5.4). t c bn phn
hu c, lm tng r rt CO
2
, c ngha
quan trng lm ngun cung cp CO
2

trong nh m trng cy.
-20 -10 0 10 4 2 1
cm
2
CO
2
-Flux.10
6
gCO
2
/cm
2
.s ft(z).10
2
cm
3

F
t
= 40
k = 0.5
Q
N
= 10ly/min
K
N
= 500cm
2
/g
Nh trn ni, mc d quan h
v tc dng ca mi mt nhn t
c nghin cu r rng trn mc
no , nhng khng th cn c vo
thc nghim ca m thc m hnh ho
nhn ra phng hng iu khin
thch ng i vi quang hp ca qun
th cy trng, vn cn phi c nhng
sa i tng ng vi m hnh theo
mc ch yu cu.
Hnh 5.4. Quan h ca s phn b nng CO
2
trong qun th cy trng vi CO
2
t t thi ra
Hnh trn: Nng CO
2
; Hnh di: Thng lng CO
2
Trng trt theo k hoch nng sut cao
Lin X (c), ngi ta tin hnh th nghim trng ng theo k hoch nng
sut cao vng Vonga. Ngi ta ln lt xt n cc ch tiu c bn ca qun th
cy trng nh sau:
1. ng cong tng din tch l
2. To thnh th nng quang hp cn thit c hiu sut quang hp nht nh
3. H s s dng nng lng mt tri trong quang hp
4. H s thot hi nc.
Vic lp trnh t ca chng da vo cc s liu di y: Vi iu kin ca vng
Vonga, t ngy 20 thng 5 ny mm n ngy 20 thng 9 thu hoch, trong thi gian
ng ht 1.800 triu kcal/ha, trong s 3.000 triu kcal/ha bc x quang hp c ti
mt t, tn tr vo trong ht ng nng lng 220 - 270 triu kcal/ha (tng ng vi
khi lng cht kh 55 - 65 tn/ha, hoc khi lng ht 20 - 25 tn/ha). l con s
kh nng ln nht ca nng sut ng thi y. C nm, ngi ta t mc tiu l t 1/2
con s , tc l thu c 12 tn/ha ng ht. Nu t l cht kh tch lu trong ht l
0,43 - 0,45 th cn c nng sut trng lng cht kh tng cng l 23 - 24 tn/ha.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

135
Trong iu kin nc trong t thch hp v phn bn , hiu sut quang hp bnh
qun trong ton thi gian sinh trng ca ng l 7- 8 g/m
2
ngy. Nu ly hiu sut
quang hp bnh qun l 8 g/m
2
ngy, th to thnh 24 tn cht kh/ha s cn th nng
quang hp (din tch l tnh ton trong c thi gian sinh trng, ly n v l hecta) l
khong 3.000 nghn m
2
ngy.
Nu ch s din tch l ca qun th ng vo thng 7 v 8 l 4,0 v 6,0 th tri qua
c thi gian sinh trng s c th hnh thnh th nng quang hp ni trn.













5.5
35
125
172
395
345
370
298
196
204
152
0
20
40
80
2
4
1
3
A
5,6
34
148
282
180
370
400
380
230
146
200
2
4
1
3
0
5
10
15
20
25
30
35
B
0
Thng 6 7 8 9 Thng 6 7 8 9
Hnh 6.4. Tr s o thc ca trnh t th nghim trng trt theo k hoch
i vi ging ng VIR-156
A: trnh t, B: tr s do thc (1964)
1. Din tch l x 10
3
m
2
/ha; 2. Hiu sut quang hp (g/m
2
ngy); 3. Lng tng trong ngy
ca cht kh (kg/ha); 4. Khi lng cht kh ton b (tn/ha)
14
12
10
8
6
4
2

300 8
8
23
43
56
60 59
55
51
45
32
1
3
1
A
2
7
25
48
43
61
56
53
51 45
33
1
B
2
1
2

6
200


4

100
2

0 0



Thng 6 7 8 9 Thng 6 7 8 9
Hnh 7.4. Lng thot hi nc v h s thot hi nc ca ng trng trt
theo k hoch ging ng VIR-156
A: trnh t, B: tr s do thc
1. Lng thot hi nc (tn/ha.ngy); 2. Lng nc tiu hao cho thot hi nc (x10
3
tn/ha);
3. H s thot hi nc;

Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Da vo cc s liu ni trn, c th tm thy c ng cong trnh t sinh trng
din tch l 1 v ng cong trnh t hiu sut quang hp 2 ca ging ng VIR - 156
(hnh 6.4.). c ng cong din tch l v quang hp, tri qua c thi gian sinh
trng s c th tm c lng tng khi lng cht kh ca tng 10 ngy mt (hnh
6.4., ng cong 4). Nng sut ton b cht kh cui cng l 24,3 tn/ha. ng thi,
cn c vo chnh lch gia tr s t sut nng lng mt tri v nng lng mt tri
c ht l c th tm c nng lng tiu dng cho thot hi nc (hnh 7.4.).
Lp k hoch bn phn v k hoch ti nc: t ch s din tch l s ln nht
ca VIR-156 l 4,0, th nng quan hp l 2927 nghn m
2
ngy, tr s thc t tng ng
l 3,7 v 2908 nghn/m
2
ngy. Tr s k hoch v tr s o thc ca hiu sut quang hp
bnh qun ln lt l 8,3 g/m
2
ngy v 8,3 g/m
2
.ngy; tr s o thc l 23,8 tn/ha. Tr
s bnh qun trong tun (10 ngy) ca hiu sut quang hp o thc v tr s thit k
chnh lch nhau rt r rt. Nguyn nhn l do trong thi gian hiu sut quang hp gim
thp, bc x quang hp c tng i t. H s s dng hu hiu quang hp bng tr
s k hoch l 0,45.
Kt qu trn chng t, thng qua iu khin quang hp s c th thc hin c
con ng t nng sut cao ph hp vi h s s dng nng lng bc x mt tri theo
k hoch nh trc. Trng im ca k hoch ny l ch lm th no duy tr c
quang hp mnh, v th k hoch bn phn v ti nc l rt quan trng. lm cho
vic sn xut cy trng tr thnh h thng iu khin thch ng, phi c trnh t ho nh
ni trn, m m thc ho h thng tr nn rt quan trng.
S vic nu trn l mt th d thnh cng, thc t khi thc hin k hoch nh vy
c th xut hin rt nhiu vn , trong vic d bo kh tng s l mt vn rt
quan trng.













136
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

TM TT
H sinh thi ng rung trong iu kin hot ng ca con ngi, khng
ngng c iu chnh theo chiu hng ngc li vi quy lut t nhin
cung cp nhng sn phm cn thit cho loi ngi. Phng php iu khin
c th chia ra lm ba kiu: iu khin vt l, iu khin ho hc v iu khin
sinh hc. Trong chng ny cp n cc kha cnh ca h thng iu
khin thch ng v ng dng cc m hnh trong iu khin qun th cy trng.
iu khin v ging v nng cao hiu sut quang hp l nhn t quyt nh
trong vic nng cao nng sut ca h sinh thi ng rung. ng thi iu
khin nhit thng qua nh knh ngy mt tr nn ph bin cc nc vng
n i v nhit i. Bn thn nh knh c th c coi l mt h sinh thi ng
rung trong cc thng s v nhit c con ngi kim sot mt cch
tch cc.


CU HI N TP
1. Anh/ch hiu nh th no l iu khin h sinh thi ng rung?
2. Trong thc tin sn xut Vit Nam, ngi nng dn iu khin h sinh thi
rung la nc nh th no?
3. Hy nu cc u im ca vic p dng m hnh ha trong iu khin h sinh
thi ng rung?
4. Anh ch hy nh gi kinh nghim truyn thng ca ngi nng dn Vit
Nam thng qua cu tc ng sau Nht nc, nh phn, tam cn, t ging.
Th t ca cc bin php ny thay i nh th no trong iu kin hin
nay? Hy chng minh?
5. ng cha ta thng ni ngi p v la, la tt v phn. ng v gc cc
nh trng trt, anh/ch hiu nh th no?
6. Hm s quang hp ca l ph thuc vo nhng yu t no?
7. Cn lm g nng cao kh nng quang hp ca qun th cy trng? Gii
thch?
8. Hy nu mt s v d v kh nng sng to ca ngi nng dn Vit Nam
trong qu trnh iu khin h sinh thi ng rung?

137
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

TI LIU C THM
1. Ota Keizaburo, Tanaka Ichir, Udagawa Taketoshi v Munekate Ken. Sinh thi
hc ng rung . NXB Nng nghip, H Ni 1981 (Bn dch ca on Minh
Khang).
2. Ahuja, L.R., Ma, L., and Howell, T.A. Whole System Integration and
Modeling - Essential to Agricultural Science and Technology in the 21
st

Century; pp. 1-8. In: L.R. Ahuja, L. Ma, T.A. Howell (eds.), Agricultural
system models in field research and technology transfer. Lewis publishers;
Boca Raton, USA. 2002.
3. Nguyn Cng Tn, Ng Th Dn, Hong Tuyt Minh, Nguyn Th Trm,
Nguyn Tri Hoan, Quch Ngc n. La lai Vit Nam. NXB Nng nghip.
H Ni. 2002.
4. Nguyn Th Trm. Kt qu chn to ging la lai 2 dng TH3-3 vi thi gian
sinh trng ngn, nng sut v cht lng cao. Tp ch Nng nghip v Pht
trin nng thn 6: 686-688. 2003.
5. Nguyn Tt Cnh. Nghin cu m hnh m phng ng thi m t v
chn on nhu cu ti cho ng v u tng trn t bc mu ng Anh v
ph sa sng Hng, Gia Lm. Lun n Tin s Nng nghip. HNNI, H Ni.
2000.
6. Phm Ch Thnh, Phm Tin Dng, o Chu Thu, Trn c Vin. H thng
nng nghip. NXB Nng nghip. H Ni. 1996.
7. Odum, E.P. Basic Ecology. Saunders college publishing. Tokyo. 1983.
8. Trn c Vin. Sinh thi hc Nng nghip. NXB Gio dc. H Ni. 1998.
9. Tsuji, G.Y, duToit, A., Jintrawet, A., Jones, J.W., Bowen, W.T., Ogoshi,
R.M., and Uehara, G. Benefit of models in Research and Decision support:
The IBSNAT Experience; pp: 71-90. In: L.R. Ahuja, L. Ma, T.A. Howell
(eds.), Agricultural system models in field research and technology transfer.
Lewis publishers; Boca Raton, USA. 2002.

138
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Chng V
K THUT HC H THNG CA H SINH THI NG RUNG
Ni dung
Th gii t nhin rt phc tp v a dng i hi con ngi phi c phng thc
tip cn mt cch h thng trong nghin cu h sinh thi ng rung. Qun th cy
trng pht trin trn ng rung c mi quan h cht ch khng ch iu kin kh tng,
t ai, ch nc m cn chu nh hng su sc bi cc mi quan h vi cc loi
sinh vt khc v cc iu kin kinh t x hi ca tng a phng. Ni dung c bn ca
chng ny l m hnh ha cc mi quan h trn nghin cu chc nng v cu trc
ca h sinh thi ng rung nhm cho mt cch nhn tng th v sn xut nng nghip.
Cc ni dung sau y s c cp trong chng ny:
Dnh cho sinh vin bc i hc:
Sinh thi hc v k thut hc h thng
Dnh cho sinh vin sau i hc:
Chun b ton hc m t v phn tch h sinh thi
M hnh ho my tnh
Phn tch h thng mt s m hnh sinh thi
Mc tiu
Sau khi hc xong chng ny, sinh vin cn:
Hiu c mi quan h gia sinh thi hc v k thut hc h thng
Nm c cc phng php m t v phn tch h sinh thi
Hiu c cch thc x l m hnh sinh thi trn my tnh (sinh vin sau i hc)
1. Sinh thi hc v k thut hc h thng
Tnh tng hp ca sinh thi hc
Nghin cu khoa hc thng c hai hng chnh: mt l c gng phn chia i
tng nghin cu thnh nhng phn rt nh, rt thun; hai l hng tng hp t chc
nhng i tng chia nh li. Phng php lun ca hng th nht l rt ly mt phn
t trong h thng thc t ht sc phc tp, c gng c lp n vi mi trng xung
quanh, cu thnh mt trng thun nhit v m c nh c li cho thc nghim,
tm ra quy lut no trong phn h thng ; trnh nhng ci bn ngoi h thng c
nghin cu ln vo trong phm vi thc nghim, tm mi cch lm cho h thng thc
nghim tr thnh thun khit nht v thun tu; thm ch ph hoi c m t bo phc
tp, lm i lm li ly ra mt loi men no , ri dng men thun tin hnh
thc nghim sinh ho theo kiu h thng ng nghim.
139
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Mt h thng thc nghim d l thun n u, nhng nu nghin cu t m hn,
th h thng li c th c cu thnh bi nhiu thnh phn th cp, ngha l vic
chia nh li c tip tc khng gii hn.
Mt pht hin mi bt k no trong nghin cu kiu chia nh nh vy, ch cn
n c lin h vi bn cht ca cng s vt, c khi cng c hiu qu trc tip v c tnh
ng dng tng i ln. Th d: nu pht hin c mt cht no c tc dng lm
tn thng mnh i vi h thng h hp hoc h thng quang hp ca sinh vt, c th
l mt phn cc nh, s c th tr thnh mt bin php c hiu qu hn ch su bnh hi
v c di.
Cho n nay, phn ln cc nghin cu khoa hc u theo phng php chia nh
nh vy. Nhng kt qu nghin cu nh th, mt khi ng dng mt cch n thun vo
trong thc t phc tp, thng lun b va vp, c khi cn cho kt qu tri ngc vi
mun. Th d: vic phun thuc bo v thc vt nhm bo v cy trng v tnh chng
thuc ca su bnh. Loi c di t b tc dng ca thuc tr c li pht trin mnh khi ta
dng thuc tr c (nh loi Eleocharis trong rung nc). T nhng thc t , con
ngi nhn thc c rng t nhin l phc tp, do phi i x vi n nh nhng s
vt phc tp v cn phi tin hnh nghin cu tng hp. T , mt s thut ng nh
h thng, k thut hc h thng c s dng ngy mt ph bin hn.
Nh ni trn, c kh nhiu phng php phn tch m sinh thi hc p dng,
nhng suy cho cng u xoay quanh yu cu tng hp. Sinh thi hc l mn khoa hc
c tnh tng hp rt cao. Bi v: 1) s hnh thnh ca sinh thi hc cn tng i tr,
cn cha c chia nh ra; 2) sinh thi hc l mt mn khoa hc phi ly a bn
nghin cu thc a lm chnh pht trin; 3) iu kin thc t, quan h gia sinh
vt v mi trng, quan h gia sinh vt vi sinh vt rt phc tp c v cu trc v chc
nng, khng d dng g m ly mt phn a vo phng th nghim. Ngi ta ni tnh
tng hp ca sinh thi hc rt cao cng th hin nhng mt .
Trong lnh vc k thut, gn y vic trang b c gii cho sn xut tr nn v
cng phc tp v vi quy m ngy cng ln. Khi dng b phn kin trc trc y
nghin cu hot ng chnh th ca nhng trang thit b ny, do nhng chnh th ny
qu phc tp, nn sinh ra quan nim h thng (system concept). Mt s h thng
phc tp nh vy c tng hp li vi nhau v mc ch nht nh, hoc c vn
dng theo mt quy lut nht nh (phng php c tnh ph bin). Nhng phng php
tng hp ny c pht trin khng ngng v c gi l k thut hc h thng (system
engineering).
Phn sau s ni n qu trnh nghin cu ca sinh thi hc v k thut hc thot
nhn hnh nh tri ngc nhau. i tng nghin cu ca sinh thi hc tn ti t lu,
cn k thut hc h thng mi c hnh thnh. Ch ng ca hai lnh vc tuy khc
nhau, nhng khi nim h thng ca chng li ging nhau. Phn then cht ca phng
php x l h thng m k thut hc ra c ngha tham kho quan trng i vi sinh
thi hc.
140
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

141
Cu trc ca h thng
H thng: H thng bao gm nhiu thnh phn c quan h vi nhau v t hp li
vi nhau mt cch rt phc tp hp thnh mt chnh th c ngha nht nh.
Trc ht cn bn l vn xc nh cu trc ca h thng. Trong tp hp nhiu
thnh phn hp thnh, xp t ci no vi ci no vo trong mt h thng ng nhin
l c s khc nhau do mc ch nghin cu, nhng cng khng th xp t tu c.
Hnh 1.5 cho thy, gi thit c 6 thnh phn hp thnh, tp hp thnh phn [1, 2] v tp
hp [3, 4, 5, 6], nhng thnh phn trong du mc [ ] c quan h cht ch hn, v th
tr thnh cc tp hp khc nhau. Trong trng hp, nu khng c l do c bit no m
tu vch ng chm chm coi [1, 2, 3] l mt h thng th s gy kh khn cho bc
nghin cu tip theo. Ni mt cch khc, h thng l tp hp do mt s thnh phn kt
hp hu c vi nhau, c th phn bit n vi mi trng hoc h thng khc v c
tnh c lp tng i mc nht nh.

H thng 1 H thng 1I

Thnh
phn hp
thnh 1
Thnh
phn hp
thnh 2
Tn hiu
ra
Tn hiu
vo
Mi
trng
Thnh
phn hp
thnh 3
Thnh
phn hp
thnh 4
Tn hiu
vo
Thnh
phn hp
thnh 5
Thnh
phn hp
thnh 6
Mi
trng
Tn hiu
ra
Hnh 1.5. H thng l s hp thnh ca nhiu thnh phn c quan h
vi nhau, ni lin vi mi trng bng u vo v u ra












Thnh phn hp thnh: gi l thnh phn hp thnh tc l mt s b phn hp
thnh h thng, bn thn chng li do nhng thnh phn cp thp hn hp thnh.
Nhng thnh phn cp thp ny li do nhng thnh phn cp thp hn na to ra. Nh
trn ni, nu tip tc chia nh khng gii hn, th cui cng (vi trnh hin ti) c
th t n mc ht c bn. Song d khng t n mc ht c bn hay nguyn
t, chng ta cng tm hiu v nm vng h thng sinh thi ng rung, do vic
chia nh thnh phn hp thnh nn lm n mc thch hp; i vi ni dung ca thnh
phn (n cu to bi ci g) th vn phi tha nhn: c tn ti mt n v thnh phn
hp thnh m n ngi ta khng truy hi g thm na. tc l thnh phn hp
thnh m chng ta mun ni n.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

142
Thnh phn hp thnh ging nh mt chic
hp en c u vo v u ra (hnh 2.5). Ging
nh chic my t ng bn hng, b ng tin
vo (chuyn vo) th mt th hng bt ra
(chuyn ra) bt k l bao thuc l hay chai nc
qu, c cu bn trong ta nh khng suy tnh g
c. V quan h gia i lng vo v i
lng ra ca cc thnh phn c th xc nh
nh thc nghim, cng c th li dng kt qu
nghin cu ca cc nh chuyn mn lin quan.
Nhng d th no i na cng c th xut hin mt cc din nh sau, nu khng xt n
thnh phn cp thp hay cp thp hn na ca thnh phn hp thnh, s khng th biu
hin tt hnh ng ca thnh phn hp thnh phc tp hn. Trong trng hp ny,
chng ta ln lt gi l h thng con v h thng chu (subsubsystem) (hnh 3.5).
Thnh phn hp thnh Yi
Tn hiu ra
Zi [= f(Xi)]
Tn hiu vo
Xi [= f(Ym)]
Yi
(Hp en)
Hnh 2.5. S hnh khi ca thnh
phn hp thnh (yu t) ca h thng
thng thng. n v nh nht x l
coi nh hp en

Mi trng (tn hiu vo)

nh sng mt tri
Nhit khng kh
C di
Cn trng
Vi sinh vt
Thnh phn
th nhng
Vi sinh vt t

H thng ti sn xut
cht kh (cy trng)
C quan
quang hp
C quan
vn chuyn
C quan
dinh dng
C quan
lu tr
+
+
+
+








H thng
Thnh phn
hp thnh
H thng chu
thnh phn hp thnh
H thng con chu Thnh phn
hp thnh
H sinh thi ng rung



Hnh 3.5. Quan h ca thnh phn
hp thnh h thng, h thng con,
h thng chu
Hnh 4.5. Phm vi ca h thng.
C s khc nhau do phm tr vn
khc nhau m ngi ta nghin cu
Gi l thnh phn (yu t) ca h sinh thi ng rung l ch qun th cy trng, c
di, qun th cn trng, NHB
3
trong t, khi lng v s lng vi sinh vt t...
H thng v mi trng: Mi trng ca h thng l tng hp tt c cc thnh
phn bn ngoi h thng, thuc tnh ca n thay i s c nh hng n h thng v
ngc li, do hot ng ca h thng m thuc tnh ca thnh phn mi trng cng b
nh hng theo.
Trn thc t, ci c coi l yu t bn trong h thng v ci g c coi l mi
trng l do cch nhn ca con ngi i vi h thng, nht l quy m ca h thng
c m rng n mc no, di ca to thi gian xem xt (ly vn pht
sinh trong my thng lm i tng, hay xem xt thi gian sau 10 nm, 20 nm) khc
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

nhau m c nhiu sai khc. Th d, h thng ng rung, nh hnh 4.5 cho thy, nu ly
cy trng lm chnh, th ci ngoi cy trng nh nng lng mt tri, nhit khng
kh, cn trng, c di, vi sinh vt ... u l mi trng ca n. Ly cy trng lm
chnh, l gii thch ch quan ca loi ngi li dng cy trng, nu cho rng cn
trng v vi sinh vt trong s hnh thnh h sinh thi ng rung, cng quan trng ngang
vi thc vt mi ph hp thc t, th mt b phn trong mi trng li c th c a
vo trong h thng (hnh 5.5).
Trong vic nghin cu sinh thi hc ng rung tnh tng hp rt mnh, d mi
u xut pht t h thng quy m nh, nhng theo s tin trin ca vic nghin cu
(dn dn a mi trng vo trong h thng), quy m ca h thng t nhin s c xu
th m rng, thm ch cui cng tr thnh h sinh quyn.

Y
1
Y
4
Y
2
Y
3
Mi
trng
A
Mi
trng
B
Gii hn ca h thng
Hnh 5.5. Gii hn gia h thng v mi trng
3
1
2









Nhng c trng ca h sinh thi
So snh vi h thng k thut, ni chung h sinh thi c mt s c trng sau:
1/ C nhiu phn ng tc chm hn h thng k thut. So snh qu trnh sn
xut ca nh ho hc v sn xut sinh vt cn nhiu thi gian th thy s khc nhau v
cng r rng. Do , s iu khin i vi h sinh thi ng rung, c tht ng l cn
my tnh h thng tuyn tnh (Computer online system) hay khng l vn rt cn
c quan tm.
2/ Nhng thnh phn c phn ng cc k nhanh v nhng thnh phn c phn ng
rt chm cng nm trong mt h thng. Th d, qu trnh quang hp xut hin phn ng
ly giy hoc pht lm n v; bin i hnh thi do sinh trng th cn xt nhiu ngy
thng hoc nhiu nm. Ngoi ra, nh s phn gii cht hu c trong t hay qu trnh
bin i tnh cht l ho hc ca t, cn mt thi gian tng i di mi t n cn
bng i th. Do khi xt n vn bin i trong thi gian ngn, i vi nhng
thnh phn xem ra c bn n nh, vi thi gian ko di nu vn coi chng l bt di
bt dch th thng l dn n sai lch ln. i vi h thng tn ti hn hp tc phn
ng (nh s thi gian) nhanh chm khc nhau, khi tnh ton bng my tnh, cng
thng d tr thnh nguyn nhn gy ra sai s tnh ton.
143
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

3/ Bn thn cu trc ca c cu (thnh phn hp thnh) ca h thng cng c
bin i. nh my bn thn c cu trong mt thi gian nht nh khng th bin i
ln. Cho nn cu trc quan h chuyn vo, chuyn ra ca n v thnh phn hp
thnh cng khng bin i nhiu lm, cn trong h thng sinh vt li khng h c s
bo m nh vy. Thm ch, c thnh phn hp thnh hon ton khng tn ti trong
mt thi gian no , nhng sang thi gian khc li xut hin ph thm vo trong h
thng (nh s hnh thnh c quan d tr ca cy trng).
4/ Trong quan h hm s chuyn vo, chuyn ra ca thnh phn hp thnh, phn
nhiu l c c tnh bo ho v khng tuyn tnh kh r. Th d, quan h gia nng
cht dinh dng trong t v tc ht ca r; quan h gia cng chiu sng v tc
quang hp. Do , mt lot phng php v thut ton tuyn tnh pht trin t h
thng k thut hc khng th dng y nguyn nh th, em li nhiu kh khn cho
vic x l ton hc i vi h thng sinh hc.
5/ Bt k l bn trong hay bn ngoi ca h thng cng u tn ti nhiu nhn t
con ngi kh iu khin. V th, mc d to ra m hnh ton hc hay m hnh my
tnh v tin hnh thc nghim, nhng mun chng thc kt qu thu c trong h
sinh thi thc t, c khi li v cng kh khn.
Qu trnh phn tch h thng (m hnh ho v thc nghim m hnh)
H thng k thut hc l i tng mi hp thnh, cn h thng sinh thi hc ng
rung th l i tng phn tch sn c by ra trc mt. Do , mc ch v qu trnh
phn tch hai loi h thng t nhiu c s khc nhau. Nhng im chung ging nhau l:
m hnh u c tc dng quan trng.
Qu trnh ca k thut hc: Qu trnh hp thnh ca h thng k thut hc, trc
ht l t ch to mt h thng c chc nng g, cng tc l bt u t vic xem xt t m
iu kin thit k ca n. Cn c vo nhng iu kin ny lm thnh kiu dng c
th ca bn thit k, tri qua qu trnh kim nghim cc loi chi tit, cui cng hp
thnh h thng m ta yu cu.
H thng hp thnh, trc khi a vo s dng, tin hnh chy th cc iu
kin mi trng. So snh kt qu chy th vi iu kin thit k mong mun, tin hnh
tu sa nhng ch khng thch ng. ch to thnh h thng cht lng cao, phi sa
i sa li nhiu ln.
Song, trong mi trng h thng quy m ln cn u t rt ln, vic dng thc vt
tin hnh chy th cc iu kin khc nhau, hoc xem kt qu tin hnh sa
li, ngy cng kh khn c v mt kinh t v v mt k thut, thm ch khng cho
php lm nh vy. Do , trc khi ch to vt thc, phi ch to trc m hnh, cho
thc nghim m hnh lp i lp li, tin hnh gii tch trc v sa i trc cho h
thng (hnh 5.6).
Trong k thut hc h thng, vic thc nghim m hnh tr thnh phng php
quan trng ca vic gii tch h thng v hp thnh h thng. V th phng php ch
144
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

145
to m hnh, phng php thu c ngy cng nhiu thng tin nh s dng m hnh v
thc nghim m hnh c s pht trin nhy vt. Nhng phng php cng rt c
trin vng trong nghin cu sinh thi hc.









Lp
m hnh
Thc hin
m hnh
Mch din hiu chnh
2
Sai
lch
4
3

M t cc
tnh nng
c th c
ca h
thng mc
ch
1
Hp
thnh
h thng
mc ch
5
[Th tc nghin cu k thut hc]
Cch vn dng h thng
6


Vn
dng









[ Th tc nghin cusinh thi ]
Lp m
hnh
Thc hin
m hnh
Mch din hiu chnh
2
Sai
lch
4
3

S liu
iu tra
thc
nghim
1
5

H
sinh thi

iu tra
thc
nghim
h thng
thc t
Cch iu khin h sinh thi
6
S dng
h
sinh thi

Pht hin ti thc nghim mi

Hnh 6.5. i chiu qu trnh hp thnh h thng k thut hc
v qu trnh nghin cu sinh thi
Qu trnh nghin cu sinh thi ( ngha ca m hnh trong sinh thi hc): H thng
ly lm i tng nghin cu ca sinh thi hc l ci tn ti t trc, tng i t
nghin cu sng ch mi nh k thut hc. Nhng trong sinh thi hc, h thng cn
nghin cu cng phc tp, th vic gii tch h thng cng cn phi c m hnh (m
thc).
Nghin cu sinh thi trc ht l thng qua iu tra d ngoi v thc nghim d
ngoi thu c nhng s liu v ti liu v h thng i tng nghin cu. Theo s
tin trin ca nghin cu, nhng ti liu ngy cng phong ph (m t hin tng), ri
t tm ra mi lin h c ngha ch o, mi lin h bn cht v hnh ng ca h
thng, nu ra nhng quy lut c tnh ph bin. lm cho mt s nhn thc tru tng
ho nh vy (hoc gi l gi thuyt lm vic) tr thnh hin thc, th cn phi ng dng
nhng m hnh khc nhau. Nh phn sau s ni, c m hnh thu nh tru tng ng c
ln thnh sa bn, m hnh kiu mch in, m hnh ton hc v m hnh my tnh.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

i vi m hnh, ch i hi n nm chc tht tt c cu trc l lun bn cht
ca h thng i tng nghin cu l c, cn s khc nhau v b ngoi v v cht vt
liu th khng t thnh vn . Thm ch c th s dng vt liu hon ton khc vi
cht vt liu ca h thng thc t, ch cn din t rt tt c chc nng ca n, da
vo im ny c th ni n l bng chng ca s tru tng thnh cng.
Song, nu ch cho rng m hnh c th phn nh rt tt nhn thc th khng
ng. Gi l m hnh din t rt tt c cu trc l lun ca h thng thc t c
ngha l trong m hnh cng phi c, hay pht huy c chc nng l lun tng t vi
h thng i tng nghin cu. Cho nn phi ni rng, s dng m hnh l c th tin
hnh thc nghim v cu trc l lun ca m hnh, hn na loi thc nghim ny l
c th c. Loi thc nghim tin hnh bng m hnh ny gi l thc nghim m hnh
(m hnh ho ngha rng).
Nh phn sau s ni, m hnh c nhng loi hnh khc nhau. Mt trong nhng tiu
chun nh gi i vi nhng m hnh xem vic thc nghim m hnh hoc vic
o nh kt qu thc nghim c th vn ng linh hot n mc no trong cc m
hnh .
i vi kt qu ca thc nghim m hnh cn i chiu vi ti liu ghi chp cc
hin tng ca h thng vn c, cn nhc nh gi. C khi pht hin c nhiu sai
khc, nghin cu nguyn nhn sinh ra nhng sai khc v tin hnh sa li m hnh
(hnh 6.5, 4). Sai khc gia thc nghim m hnh v hin tng h thng thc t, c th
i su iu tra h thng thc t v cung cp con ng mi (hnh 6.5, 5), cn c th
pht hin li, tm ra u mi ca h thng m m hnh c (nhn thc) cha lng thy
ht. Ngi ta gi l chc nng ca phng php pht hin bng m hnh.
Nh trn ni, mc ch s dng m hnh ca k thut hc v sinh thi hc tuy t
nhiu c ch khc nhau, nhng cch thc ch to m hnh, phng php gii tch, cng
trnh tun hon ci tin m hnh u c nhiu im tng t.
Th loi v s pht trin ca m hnh: m hnh c nhiu hnh thi; t lu ngi ta
dng m hnh thu nh. Dng vt liu c cng tnh cht nh thc vt nhng kch
thc c thu nh li nh m hnh cu cng, m hnh my bay, m hnh sa bn ca
dng sng, vnh vc; ng vt thc nghim nh chut, khi s dng thay th cho ngi,
th nghim nng hc din tch 10 mPPP
2
, h thng th nghim trong chu... y c mt
tin l, kt qu ca thc nghim m hnh nhn vi mt h s l c th tr v vi
vt thc.
M hnh phi d x l hn vt thc. M hnh thu nh tin li ch c c im l
nh, nhng khng c nhiu u im.
M hnh ho cng pht trin, ngi ta cng dng cc m hnh tuy hnh thi b ngoi
hon ton khng ging vt thc, nhng chc nng li rt ging nhau v d nm vng.
gi l m hnh tng t trc tip (direct analog model). Th d, t i h thng my mc
(h thng lc n hi - trng lc), h thng chn ng ca qu tim thnh mch in gm
146
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

147
in tr, t in, pin in (hnh 7.5). Vic ch to m
hnh mch in c t do tng i cao, vic xc
nh kt qu ca m hnh cng rt gin n. Nhng
phi ch ra rng, trong qu trnh i h thng i
tng nghin cu thnh mch in l tin hnh
tru tng ho chc nng.
Phm vi c th m hnh ho ca mch in n
gin khng rng lm. Th d trong t nhin thng
xut hin nhng quan h hm s m, quan h hm
s logarit... kh tin hnh m hnh ho bng m
hnh mch in n gin. M hnh c to thnh
do li dng tnh nng vt l ca in tr v t in,
v thuc tnh vt l ca bn thn nhng chi tit ,
nn c tn ti gii hn ca kh nng m hnh ho
nht nh. Ngha l ngi ta mun thot khi s hn ch v thuc tnh vt l ca nguyn
liu cu to thnh m hnh, biu t mt cch linh hot nhng quan h l gic ca s
vt. M hnh ton hc c th tho mn c nhng yu cu .
T in
Thi gian
Vn k
Tr khng
Hnh 7.5. M hnh hiu in th
chn ng ca tim
M hnh ton hc dng nhng k hiu hon ton tru tng d m t cc loi h
thng, v c th tin hnh tng i t do nhng thc nghim trn giy. Nhng m
hnh ton hc cng c hn nht nh. Ni dung ghi trong cng thc ton hc v tt
u c th gii c bng ton hc (gii tch hc). Mc d v mt l lun coi l c th
c, nhng tn nhiu thi gian, s thc l khng th c. Cui cng xut hin m
hnh my tnh.
Phn ct li ca my tnh in t hin i cu to bi cc chi tit in t, nhng c
khc vi trng hp m hnh mch in ni trn, hot ng ca n khng phi l trc
tip li dng tnh tng t v tnh cht vt l ca nhng chi tit in t . Bn trong
my tnh in t c nhng hot ng k thut hc in t no, chng ta hu nh c
th khng cn xt n, ch cn coi n l mt loi my mc c cc chc nng tru
tng thun tu s dng l c. Do xut hin loi cng c ny, nng cao ln
nhiu trnh t do ca thc nghim m hnh (ch cn nhn ni tnh ton cc con s).
C nhng vn m ton hc mong i m cha thc hin c th nay dng my
tnh cng thc nghim c m hnh.
Nhng loi vn cn tm li gii
a/ Phn on cu trc m hnh c tt hay khng.
b/ Tnh cc thng s cha bit.
c/ D tnh tng lai ca h thng.
d/ Suy on iu kin n nh ca h thng.
e/ Nu r phng php iu khin tt nht.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

a/ v b/ l mt phn ti trong nghin cu, mc ch l gii quyt nhng nghi vn
v nhn thc (gi thuyt lm vic) ca ngi nghin cu. y mun ch rng khi
c s liu v hin tng ca h thng thc t, vn l lm th no ni dung ca hp
en (m hnh) khp vi nhng s liu . Dng ngn ng ton hc ni, tc l tng
ng vi vn bin tr v vn tr s cho sn.
c/, d/, e/ l vn sau khi lm ra c m hnh tng i ng c tin cy, vn
dng cc m hnh tm li gii. c/ l mt vn ca tr s ban u. Th d, iu kin
ban u v iu kin chuyn vo no , qu trnh sinh trng v nng sut c cy trng s
bin i th no; trong trng hp gim lng dng thuc bo v thc vt, d tnh b mt
ca h sinh thi ng rung sau 10 nm s thay i ra sao, .... d/ l vn c th nghin cu
lm th no khi phc c s cn bng ca cc h sinh thi ang lun lun b ph hoi.
Th d, i vi h thng su bnh hi d pht sinh ln c tnh chu k, lm th no dng
tng i t thuc m dp tt c, lm th no su bnh hi dng li mc tng i
thp, u thuc loi vn ny. Phng php phn on tin cy ca gii p, phng
php x l ton hc ca chn ng, phng php tnh ton ca vn tr s c sn... u c
ch i vi cc vn ni trn, e/ v d/ l vn lm th no h thng nghin cu c
tip tc vn dng theo mc ch, cng tc l ni vn lm th no tm c phng php
. i vi h thng nng nghip trang b thit b iu khin tt, tm phng php qun
l hp l l ti tiu biu cho loi vn ny.
2. Chun b ton hc m t v phn tch h sinh thi
Gn y do s pht trin ca my tnh, nht l s tin b ca thit b mm, d
khng c nhiu kin thc ton hc nhng chng ta cng c th tin hnh phn tch h
thng. giai on hin nay, phn tch ton hc vn c ngha quan trng v hai l do:
mt l khi lp m hnh my tnh, nu c kin thc ton hc th vn c li; hai l s suy
din ton hc i vi vn lm cho chng ta thu c cng nhiu thng tin quan trng
khi qut v h thng hn l gii p ca my tnh tnh ton tr s.
Cng c ton dng din t h thng
H thng v h phng trnh: Nh trn ni, h thng do nhiu thnh phn hp
thnh. Bc th nht ca m hnh ton hc ly iu tra, thc nghim v cn c l lun lm
c s, lm r nhng quan h hm s gia i lng vo v i lng ra ca mi thnh phn
hp thnh, ri biu hin thnh cng thc (n y, cha c g khc vi trng hp khng
c khi nim v h thng).
V khi nim h thng, tin ca n l tha nhn gia cc thnh phn khc nhau
ang tn ti quan h qua li, do , trong mt b phn i lng vo ca cc thnh
phn hp thnh cng bao hm i lng vo ca cc thnh phn hp thnh khc. Ni
mt cch khc, hm s biu th i lng ra ca mt thnh phn hp thnh no l ly
i lng ra ca mt thnh phn hp thnh khc lm bin s m to thnh.
By gi gi thit h thng hp thnh bi n thnh phn, i lng ra ca cc thnh
phn ln lt l y
1
, y
2
, ... y
n
th:
148
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

149





Biu thc ca cc thnh phn u do i lng chuyn ra ca cc thnh phn khc
cu thnh, nn khng th tm gii tng biu thc ring, m lm nh th cng khng c
ngha. Mun tm y
1
, phi bit y
2
, mun tm y
2
phi bit y
1
. Nh vy l xut hin mt
trng thi quan h qua li nhn qu. trng thi ny, lm r hnh vi ca tng th ca
chng, cn suy din vi iu kin ng thi tho mn cc h thc, cng tc l gii bi
ton vi h n phng trnh. Do , im quan trng u tin m ton hc v h thng
biu th l: lp mt h phng trnh vi s lng thnh phn hp thnh h thng.
Trng thi n nh v trng thi qu : Trong
thnh phn hp thnh (hoc h thng tp hp ca n)
c hai trng thi quan trng: trng thi n nh v
trng thi qu . Nu i lng vo ca thnh phn
hoc h thng bin i nhanh sang trnh mi, th
i lng ra cn tri qua cc giai on qu (hnh
8.5) mi t n c trng thi n nh mi. Nh
ci ngt in, ch trong nhy mt c th lm bin i
u vo, bc vo trng thi n nh mi (hnh 8.5,
1). Nhng nh th thng tng i t, phn nhiu l
(nht l h thng cu to phc tp) t nhiu vn c
mt thi gian chm sau, tc l trng thi qu c bin s ph thuc thi gian ri sau
mi t n trng thi n nh mi (hnh 8.5, 2, 4).
y
1
= f
1
(y
1
, y
2
, ... y
n
)
y
2
= f
2
(y
1
, y
2
, ... y
n
)
.
.
.
y
n
= f
n
(y
1
, y
2
, ... y
n
)

(1)

Thi gian (t)
5
4
3
2
1
Hnh 8.5. Nhng loi hnh phn
ng qu khc nhau xy ra khi
mc chuyn vo t1 bin i
thnh dng bc thang
Biu hin trng thi qu (phng trnh vi phn). Khi chng ta nghin cu hot
ng ca h thng, c trng hp coi s bin i qu (thi gian dch chuyn) l mt
vn nghin cu; c trng hp ch hn ch vn trng thi n nh. M hnh
loi trn gi l m hnh ng; thng biu th bng h phng trnh vi phn hay phng
sai phn nh sau:
dy
1
df
= f
1
(y
1
, y
2
, ... y
n
, k
1
, k
2
, ... k
m
)
dy
2
dt
= f
2
(y
1
, y
2
, ... y
n
, k
1
, k
2
, ... k
m
)
.
.
.
dy
n
dt
= f
n
(y
1
, y
2
, ... y
n
, k
1
, k
2
, ... k
m
)
(2)







Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

150
Cch gii h phng trnh ny l tin hnh tch phn, ln lt tch cc thnh phn
hp thnh cng thc ch do thng s (k) v thi gian (t) cu thnh (hm s hin), tc l:





ngha din t ca phng trnh vi phn: Xut hin trong h thng thc t khng
phi l lng vi phn (tc ), m l lng tch phn (lng hin ti), ci cui cng
mun tm hiu l phng trnh ca lng tch phn c lin quan, nhng ti sao li u
dng phng trnh vi phn biu th trng thi ca h thng ?
By gi c lng y no (th d s c th ca mt loi no ), lng tng thm
v cht i u thnh t l y (th d t sut tng thm l a, t sut cht i l b), trng thi
nh vy dng hnh thc vi phn vit l:
dy
dt
= ay- by (4)
Tin hnh tch phn (4) th tr thnh hnh thc biu hin tch phn ca cng trng
thi :
y = y
0
e
(a-b)t
(5)
Ni chung, cui cng chng ta mun tm hiu l (5), cng thc biu th c lng
tch phn (lng hin ti y) bin i nh th no theo thi gian, ng nhin cng phi
xem l vn nh th no. Nu l h thng tng i phc tp, mun trc tip c
c hnh thc tch phn nh cng thc (5) t trng thi quan h m t bng ngn
ng ni trn l vic rt kh khn. l l do th nht phi trc ht xut pht t
phng trnh vi phn.
Nh th, c phi c ngha l cng thc (4) ch l
mt qu trnh hay cc bc cn thit dn n cng
thc (5) hay khng? khng, nht quyt khng phi
nh vy. Tc (lng vi phn) d phi hay khng
phi l s thc o, cng u c tc dng kt hp thnh
phn hp thnh (lng tch phn) vi thnh phn hp
thnh, v lm n v biu hin quan h gia cc thnh
phn, l mt lng c ngha quan trng. Quan h
gia nguyn nhn v kt qu biu th nh sau: ...
lng tch phn (lng vi phn) lng tch phn
(lng vi phn) lng tch phn ..., lng tch
phn ny nh hng ti mt lng tch phn khc
y
1
= f
1
(k
1
, k
2
, ... k
m
, t)
y
2
= f
2
(k
1
, k
2
, ... k
m, t
)
.
.
.
y
n
= f
n
(k
1
, k
2
, ... k
m,
t)

(3)

Y
Y
by
t ) b a (
0
e y y
by ay
dt
dy

=
=
Thi gian (t)
ay
Hnh 9.5. Hnh thc vi phn
v hnh thc tch phn ca m
hnh ly s tng thm s c th
gin n l v d
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

151
cch mt lng vi phn. Ch c dng phng trnh trng thi (phng trnh vi phn)
lin h tc mi biu hin c trc quan r rng quy lut lin h bn cht gia cc s
vt qua thi gian (quy lut hnh ng). l l do th hai dng hnh thc vi phn
(phng trnh vi phn) biu hin trng thi quan h h thng.
trn c dng t qua thi gian, ng nhin thng s ca phng trnh bin i
theo thi gian. Nhng cu trc logic chnh th biu th bng hnh thc vi phn (nu n l
bn cht) th khng th d dng bin i theo thi gian (hnh 9.5).
Din t trng thi n
nh (phng trnh i s):
Gi l trng thi n nh tc l
trng thi cn bng t n
sau khi kt thc trng thi qu
gy nn do bin vo (hoc
thng s h thng). Trn thc
t, trng thi n nh cng c
th hiu l mt trng hp c
th trong trng thi qu .
Nu khng im ny, c
khi ch xem xt quan h
chuyn vo bin ra trng thi
n nh ca mt h thng
(hoc thnh phn hp thnh) no . Th d, ng cong cng nh sng - quang
hp (hnh 10.5, b) m chng ta tm c qua thc nghim, quan st t m nh hnh (a)
cho thy, l s hp hp thnh ca tng phn (tr s n nh) kt qu ca nhng bin
i qu . Dng ngn ng ton hc ni, gi l trng thi n nh tc l trng thi
m h s vi phn ca m hnh ng (phng trnh vi phn) bng khng (h thng khng
c trng thi n nh, chng hn nh trng hp bin vo hay thng s l bin s thi
gian... khng thuc loi ny). Ni cch khc, din t trng thi n nh ca h thng l
trong cng thc (2) vi l h phng trnh i s din t h thng.








dy
1
dy
2
dt
= 0,
dt
= 0 th
0 = f
1
(y
1
, y
2
, ... y
n
, k
1
, k
2
, ... k
m
)
0 = f
2
(y
1
, y
2
, ... y
n
, k
1
, k
2
, ... k
m
)
...
0 = f
n
(y
1
, y
2
, ... y
n
, k
1
, k
2
, ... k
m
)
(6)
Hnh 10.5. Bin i qu (a) khi bin vo tng
thnh dng bc thang. Quan h bin vo bin ra (b)
ch trng thi n nh biu th theo kt qu ca (a)
b) a)
Lng chuyn vo Thi gian
Chuyn ra
Chuyn vo

y khng gm bt k bin s thi gian no, cho nn gi l m hnh tnh. Mt s
trng hp ni n di y u c vn ca m hnh tnh. Chng hn nh mun bit
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

152
quan h trc tip ca iu kin hin ti (iu kin ban u) vi trng thi tng lai m
trng thi n nh d tnh; hay nh mun bit trc c nu thuc tr c c dng nh
hin nay hoc n y chm dt, th h sinh thi ng rung 10 nm sau s tr thnh
trng thi nh th no. Ngoi ra, tc p ng ca mt h thng con no bn trong
h thng nhanh hn nhiu so vi h thng con khc hay thnh phn hp thnh khc th
li t ngay n trng thi n nh. Trong trng hp ny, ch c b phn (h thng
con c tc p ng nhanh) din t bng phng trnh i s (m hnh tnh).
Phn loi phng trnh vi phn v c tnh ca h thng: C th phn loi phng
trnh vi phn theo nhng quan im khc nhau, y s bn n nhng s phn loi
c lin h th no vi cu trc v c tnh ca h thng.
Tuyn tnh v khng tuyn tnh: Ton b nhng s hng c quan h t hm s (y)
v o hm ca n (dy/dt) u l bc nht th gi l phng trnh vi phn tuyn tnh,
ngoi ra l nhng phng trnh vi phn khng tuyn tnh. Nhng h thc trong h sinh
thi, rt nhiu l khng tuyn tnh. Nh phn sau s trnh by, phng trnh vi phn
khng tuyn tnh, tr trng hp c bit ra, u ht sc kh gii. Khng nhng th, v
mt c tnh cng khc vi loi tuyn tnh (xem trang 139).
Bin s c lp: H thng m chng ta nghin cu thng coi thi gian l bin s
c lp, cn c mt bin s c lp khc. Phng trnh nh vy gi l phng trnh vi
phn thng. Th d:
dy
1
dy
2
dt
+ y
1
dt
= f (t)
Nu c 2 bin s c lp tr ln, th d ngoi thi gian ra, vn cn cp n
khng gian, th gi l phng trnh vi phn ring. Nh:

x
y
t
+

x
= f (x,y,t)
Trong khi nghin cu cu trc nhn nh sng ca qun th bin i theo thi gian,
s di chuyn vt cht gia cc c quan cu trc khc nhau, phng trnh biu din l
phng trnh vi phn ring.
Hng ca phng trnh vi phn: s bc vi phn cao nht c trong phng trnh l
hng ca phng trnh vi phn ; vi phn bc cao nht
n
n
dt
y d
th phng trnh l
phng trnh vi phn hng n. Trong h sinh thi, ni chung dng phng trnh vi phn
hng 1 l c th din t y , hu nh khng th xut hin hng 2 tr ln nh gia
tc

2
2
dt
y d
. im ny khc vi h thng c hc. Trong qu trnh gii h n phng trnh
vi phn hng 1, dng phng php kh kh mt cc bin s ph thuc khc, c
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

phng trnh vi phn ca bin s ph thuc, kt qu cng l h phng trnh hng n,
do v k thut tnh ton phi thnh thuc phng trnh vi phn hng cao.
H s c phi l hm s thi gian khng? H s khng c quan h vi thi gian
(bin s c lp), l nht nh, gi l phng trnh vi phn c h s khng i. Nu
cng bin i vi thi gian, th gi l phng trnh vi phn h s bin thin.
Khi h s l hng s th d tnh ton. Trong h sinh thi, gi tr h s phn nhiu l
bin i theo thi gian. Th d, t l phn phi sn phm trong h thng sinh trng cy
trng (h thng sn xut vt cht) c bin i rt ln theo s sinh trng, nhng c mt
s h s d c coi l bin s thi gian, mt khi lm r c ch hnh ng ca
chng, t n giai on biu hin bng thnh phn bc thp hn, s c th biu hin
tng i nhiu bng hng s khng quan h vi thi gian mi. Cng tc l mt khi
xut hin h s bin i, th cn thit phi hoi nghi: c phi l mt chng c
nhn thc khng y v cu trc logic ca h thng chng.
Bn ngoi chuyn vo c phi lm hm s thi gian khng? y l mt trng hp
c bit ca h s bin i, trong cc h s, h s ch c bn ngoi chuyn vo c
phn loi theo quan im c bin i theo thi gian hay khng. Bn ngoi chuyn vo
khng c quan h vi thi gian th gi l h thng tnm, tri li th gi l h thng
khng tnm.
dy
dt
= y (f, a) ... h thng tnm
dy
dt
= y (f, a) g (t, a) ... h thng khng tnm
Trong c th sinh vt v h sinh thi c rt nhiu hin tng chu k. Hin tng
chu k trong h thng tnm (hon cnh nht nh) gi l dao ng t kch. Hin
tng chu k ca h thng khng tnm - th d dao ng do i lng vo ca chu k
c lin h vi s t quay ca qu t - th gi l dao ng cng bc.
Ton hc ca h thng tuyn tnh
X l ton hc ca h thng khng tuyn tnh, s ni sau, kh hn nhiu so vi h
thng tuyn tnh. Nu ni vic x l ton hc (gii tch) ch pht huy c th mnh
trng hp h thng tuyn tnh, cng khng phi l qu ng. Khi cn thit phi x l
ton hc i vi h thng khng tuyn tnh, ngi ta cng thng c gng a v gn
vi h thng tuyn tnh ri mi tin hnh gii tch. Do , hng chnh ca vic gii
tch ton hc ca h thng, d sao cng phi ly ton hc ca h thng tuyn tnh (ton
hc tuyn tnh) lm ht nhn. H thng tuyn tnh c th chia ra lm hai loi: tnh v
ng. ngh xem chi tit cc sch chuyn v ton, y ch ni vn tt v h thng
tuyn tnh ng, cng tc l cch gii ca phng trnh vi phn tuyn tnh.
153
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Cch tnh c in ca phng trnh vi phn: i vi h n phng trnh vi phn
hng 1, trc tin dng phng php ton (phng php thay k hiu d/dt l D tnh
ton) a v h phng trnh i s tuyn tnh tnh ton, kt qu l ch c mt bin
s ph thuc no (y
1
). Cn D c dn ti c n phng trnh i s bc n (phng
trnh vi phn hng n i vi y). Mt trong nhng phng trnh c dng chung l:
a
n
D
n
y + a
n-1
D
n-1
y + ... + a
1
Dy + a
0
y = F (t) (7)
C s ca cch tnh c in ny c xy dng trn cch tnh thng thng ca
phng trnh (7), coi l tng ca hai phn sau y tnh ton
y (t) = y
c
(t) + y
p
(t) (8)
y
c
(t) gi l nghim c bn, do c cu bn trong ca h thng quyt nh, l phn
khng c quan h vi i lng chuyn vo F (t) ca h thng. Khi t , h thng
trng thi n nh, phn ny s khng cn na, y
p
(t) gi l nghim ring, l phn c
nhiu bin i do dng hm s i lng vo (sng xung kch, sng hnh sin,...). Trong
phng trnh (7), gi thit khng c i lng chuyn vo, tc l khi F (t) = 0 v tm
nghim s tng qut ca phng trnh thun nht c th tm ra nghim c bn y
c
(t).
a
n
D
n
y + a
n-1
D
n-1
y + ... + a
1
Dy + a
0
y = 0 (9)
tnh ton (9) cn c mt phng trnh c trng (mt loi phng trnh ph) l:
a
n
r
n
+ a
n
r
n-1
+ ... a
1
r + a
0
= 0 (10)
Tm nghim r ca 10, nghim ca phng trnh l r
1
, r
2
, r
n
... th cch tnh ca
phng trnh (7) l:
(11)
t
n 1 n
t
2
t
1
r
n
t r
1 n
r
2
r
1 c
r C e C ... r C c C ) t ( y + + + + =

Nu trong cc nghim s r
1
... r
n
ni trn c my nghim s bng nhau (chng hn
trong s n nghim s c k nghim bng r
1
), i vi nhm cc nghim s bng nhau ny,
phng trnh (11) gm c cc s hng vit gp li, tr thnh dng thc sau:
(C
1
+ C
2
t+ ... + C
k
t
k-1
) e
nt
(12)
Nh trn ni, im then cht ca phng php tnh ton h thng khng n nh
tuyn tnh l vic tnh ton nh th no i vi phng trnh c trng (phng trnh
i s) bc n vi phng trnh (10). Phng trnh c trng l bc 2, th tnh ton bng
cng thc tnh nghim ca phng trnh bc 2 quen thuc. Trng hp l bc 3 th dng
cng thc Cardano, khi bc 4, vn c th tnh c bng phng php Ferrari, nhng
cng kh tn cng sc, m p n tm ra ht sc phc tp, tnh trc quan tng i
mng manh. Do m hnh ton hc c l tc dng pht hin trong nghin cu sinh
thi hc nhiu khi cng kh m thc hin. Thc t cho thy, trng hp bc 5 tr ln
khng th tnh ton c bng cng thc gii tch. Cho nn i vi h thng bc cao
phi dng phng php tnh s. Trong phng php tnh s, thng s h thng ca
phng trnh (7) khng c dng k hiu a
0
, a
1
, .... a
n
, phi dng tr s c th. V th,
quan h bnh thng gia a
0
v nghim (y) nh th no, s khng th trc tip tnh ra
154
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

155
c. Vic tnh ton nhng tr s ny thng thng phi nh my tnh in t. Khi tn
hiu chuyn vo ca h thng F(t) 0 (gi l phng trnh khng thun nht), trong
tnh ton thng thng s c thm y
p
(t). i vi hnh thc F(t) bt k cng u c
phng php chung tm y
p
(t), nu i lng chuyn vo F(t) l cc dng hm s
trong bng 18 hoc tng ca chng, dng phng php thay th h s l c th gii
c d dng. Nu ly tin l F(t) ca bng 18 v y
p
(t) tng ng (lc b phn
chng minh) th trong phng trnh (7) y = y
p
(t). Ly y
p
(t) thay vo y ca (7), lm cho
h s cc s hng ca hai v tri v phi bng nhau v xc nh h s B
0
, B
1
, B
2
, ... t
m tm c nghim y
p
(t).
Di y l tnh ton ca mt th d n gin:
th d: Th gii D
2
y + aDy + by = ct. Trc tin gii phng trnh thun nht.

+
= t
2
b 4 a a
c t
2
b 4 a a
c yc
2
exp 2
2
exp 1

tm ra nghim y
p
cn tra trong bng 21 tm ra y
c
tng ng vi F(t) = ct. Tm
thy trong bng y
c
= B
0
t + B
1
, thay n vo y ca phng trnh u c:
bB
0
t + (aB
0
+ bB
1
) = ct
do tm ra:

b
c
B
0
=

B
1
= - ac
Vy cch gii chung ca u nh sau:
ac t
b
c
t
2
b 4 a a
c t
2
b 4 a a
c y
2
exp 2
2
exp 1
+

+
=

Phng php gii bin i Laplace: Nu dng phng php bin i Laplace
gii mt phng trnh vi phn hng n so vi phng php c in ni trn, cng khng
c u im g c bit. Tm li c hai phng php u khng th khng gii phng
trnh i s bc n, v im ny m ni th khng c g khc nhau. Nhng khi i tng
l h thng hp thnh bi nhiu bin s (h phng trnh vi phn) th phng php bin
i Laplace li c mt s u im. Dng phng php ny c th din t c hai phn
ca h thng, phn ton b iu kin ban u v iu kin chuyn vo, v phn c tnh
vn c bn trong h thng (hm s truyn t), din t ring tng phn. Ngi ta cho
rng, l mt trong nhng nguyn nhn thnh hnh ca phng php bin i Laplace,
phng php hm s truyn t ly lm c s v phng php biu trong cng tc
iu khin k thut v phn tch h thng.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

156
Bng 1.5. Quan h gia hm s chuyn vo h thng v dng gii ring
Hm s chuyn vo h thng F (t) Dng gii ring y
p
(t)
nh s b B
M bt
n
B
0
t
n
+ B
1
t
n-1
+ ... + B
n-1
t

+ B
n
Lu tha thc be
kt
Be
kt
Sin bsin
f
t

B
0
cos
f
t + B
1
sin
f
t
Cosin bcos
f
t B
0
cos
f
t + B
1
sin
f
t
Cch gii phng trnh vi phn tuyn tnh bng phng php bin i Laplace gm
ba bc sau:
1. Bin i phng tnh vi phn bin s thc thnh phng trnh i s bin s
phc s (bin i Laplace thun, L).
2. dng s tm nghim, tin hnh x l i s i vi phng trnh ca n (ngha
l lm thnh dng L [y] = f (s). d tin hnh qu trnh bin i nghch bc sau,
phi thay i dng ca f(s) (s dng nguyn l khai trin phn s).
3. Bin i dng thc trn thnh lnh vc thi gian thc t (t) v tm ra nghim
(bin i Laplace nghch (L
-1
). C th dng bng bin i Laplace tnh sn bin i
Laplace thun v nghch i vi nhng hm s thi gian v c tnh tiu biu thng
xut hin (chi tit bng ny cc sch gio khoa v iu khin hc, trch ra mt phn
nh bng 1.5).
Bng 2.5. Bng bin i Laplace ca hm s thi gian
Hm s thi gian f(x) Hm s F(s)
Hm s step U
-1
(t)
s
1

t
2
1
s

be
-at
a s
b
+

) 1 (
1
at
e
a

) (
1
a s s +

a b
e e
bt at



) )( (
1
b s a s + +

sin

t
2 2

+ s

cos

t
2 2
+ s
s

Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

157
Ngoi hm s thi gian ra, nhng bin i tnh ton vi phn cng l cn thit, lit
k nh sau:
L[y] = Y
y sY
dt
dy
L =


0
0
2
2
2
dt
dy
sy Y s
dt
y d
L


0
1 n
1 n
,... 0
2 n
0
1 n n
n
n
dt
d
dt
dy
s y s Y s
dt
y d
L



y, y = f(t), Y = f(s), ch s 0 nh bn cnh [ ] biu th iu kin ban u.
Trn y, ni nhng iu kin cn thit ti thiu ca s bin i Laplace, di
y ly mt th d ng dng phng php .
th d:
t
e y
dt
dy
= + trong t = 0 , y = 3.
Bin i Laplace thun i vi c hai v
] e [ L y
dt
dy
L
t
=

+ (13)

1 s
1
Y 3 sY

= + (14)
Tm Y t h thc (14), tin bin i Laplace, v phi khai trin phn s tr
thnh

) 1 s )( 1 s (
2 s 3
Y
+

= (15)

1 s
2 / 1
1 s
2 / 5

+
+
= (16)
Tin hnh bin i Laplace nghch theo bng 19, bin thnh hm s thi gian thc
t, tc l tm c nghim:

+
=

1 s
2 / 1
L
1 s
2 / 5
L ] Y [ L
2 1 1
(17)
Vy
t t
e
2
1
e
2
5
y + =

(18)
Ton hc cc h khng tuyn tnh
Nh trc trnh by, cho n hin nay vn cha c phng php chung tm
nghim ton hc cc h khng tuyn tnh. Nhng khng phi l khng c bin php,
chng hn nh lm cho n gn vi tuyn tnh hoc tip cn bn cnh.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

158
Trng hp kt hp tuyn tnh thng s cha bit: xc nh c phi l tuyn
tnh hay khng, thng ch yu l xt n bin s. Khi nh tr s c th o c ca
bin s suy on thng s cha bit. Th d c mt m hnh tnh khng tuyn tnh
no , biu th nh h thc sau:
...
1
0
2 3 2
2 2
1
+ + +
+
= x a y a
y x
a
z
(19)
Bin s ca h thng x, y, z c th o c; do , dng my tr s o lp i lp li
tm thng s cha bit a
1
, a
2
, a
3
... trn thc t ch l mt vn gii h phng trnh
bc nht.
Phng php xp x tuyn tnh: Phm vi bin i ca i lng vo ca h thng
hay thnh phn hp thnh ca n nh, m li trng hp ht sc tip cn vi im cn
bng hay im ng tc c bin ng, th c th lm cho h thng khng tuyn tnh
cng gn vi m hnh tuyn tnh tm nghim. Phng php gn ng tuyn tnh
thng thng l khai trin Taylor. Phng php ny c th b qua khng tnh cc s
hng bc 2 tr ln ca b phn bin i l gin n nht.
th d: Trong mt h thng mng nc, cung cp nc t trc trn v thot
nc ra t l pha di, quan h gia lu lng cung cp a v mc mt nc y ca mng
nc s nh th no? H thng nh vy cng c c ng dng vo m hnh h thng
phn ng sinh ho hc, nhng gia hai loi c im khc nhau, quan h gia tc chy
ra v t mng nc v mc mt nc y ca mng nc (tng ng vi nng cht
mi trng ca h thng sinh ho hc) khng phi l t l bc 1, m l khng tuyn tnh.
t gia tc trng trng l g, theo nh l Torixenli, tc chy ra phi l:
gv 2 v = (20)
By gii din tch l thot nc l s, din tch mt ct mng nc l A, quan h
gia mc nc y vi lng cung cp nc v lng thot nc nh sau:
gy 2 s a
dt
dy
A = (21)
Do c s hng y , nn l phng trnh vi phn khng tuyn tnh. Nu mng
nc c chiu cao cao, h thng ny ri s t n cn bng. gn im cn bng
ny, quan h gia lng bin i cc nh ca i lng vo a v bin i mc nc
y c th dng phng php sau y sau khi bin n thnh gn vi tuyn tnh ri
mi gii. t mc nc ca im cn bng l y
0
, lng cung cp nc l a
0
, th
a (t) = a(t) - a
0
y (t) = g(t) - y
0

(22)
Thay vo phng trnh (21) ta c:
a a
y
y
1 gy 2 s
dt
y d
A
0
0
+ =


+ +

(23)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

159
i vi y, tin hnh khai trin Taylor i vi b phn khng tuyn tnh, th:
...
y
y
8
1
y
y
2
1
1
y
y
1
2
0 0
a
1
0
+


+ =


+ (24)
0
y 2
y
1

+ = (25)


0
y
y
Khi nh, i vi y m ni, dng hng thc bc 1 l t c gn ng.
Vy phng trnh c (21) c tuyn tnh ho v tm ra phng trnh vi phn tuyn tnh
ca b phn bin i nh sau:
a y
y 2
g
s
dt
y d
A
0
= +

(26)
Phng php nh th, trong h thng k thut hc, trng hp m i lng
chuyn vo ca n hon ton iu khin c, l c hiu qu, cn trong h sinh thi th
khng c s m bo nh vy. Ngoi ra, trong h sinh thi thc nghim khi quan st
phn ng ca chng trng hp i lng chuyn vo bin i vi phm vi ln,
phng php ny cng khng c tc dng g.
Hin tng vn c ca h thng khng tuyn tnh: Trong din bin ca h thng
khng tuyn tnh tn ti nhng hin tng c bit m h thng tuyn tnh khng c.
Vic nghin cu nhng hin tng c bit ny s gip cho vic suy on mt h thng
hay mt thnh phn hp thnh no c c ch khng tuyn tnh hay khng.
Trong phng php gii tch ca h thng, c mt phng php gi l phng
php ng p s sng chu k. Tc l i lng vo cho h thng bng i lng vo c
tnh chu k, thu c cc thng tin theo kt qu quan st chuyn ra tng ng. H thng
khng tuyn tnh lc ny: 1/ C khi trong i lng ra ca n c th gm c thnh phn
sng chu k m trong hnh sng ca i lng vo khng c; 2/ D trng hp s
sng chu k vo bin i lin tc, thu c v gc pha c khi mt im no thnh
bin i khng lin tc; 3/ Tnh n nh ca h thng trng hp tuyn tnh ch chu
s quyt nh ca kt cu bn trong h thng, cn trng hp khng tuyn tnh th
chu s nh hng ca lng to nh, hnh dng v iu kin ban u ca i lng vo;
4/ trong mt h thng dao ng khng tuyn tnh no , do s thay i ca i lng
vo, d bin v chu k dao ng c mt lc no b ri lon, nhng vn c mt s
sng c tnh cht khi phc chu k v bin c (gi l chu k gii hn). C ngi
kin ngh (Munekata, Koga, 1971), m hnh s sng chu k nh im 4/ c th coi l
m hnh ca hin tng chu k ngy ca sinh vt hay khng. Nh mi ngi u bit,
hin tng chu k ngy ca sinh vt c tnh n nh chu k tng i cao i vi mi
s bin i ca hon cnh.
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

160
Ton hc ca hin tng chu k
Gii sinh vt c kiu loi hin tng chu k, th d hin tng pht sinh ln
chu k ca cn trng v ng vt nh; cc loi ng vt v thc vt c iu kin nht
nh c th biu hin hin tng chu k tnh cht 24 gi; tnh chu k bin i hng lu
ng nguyn sinh cht (chu k 2 - 5 pht)... y, vic chun b ton hc biu din
v gii tch i vi nhng hin tng chu k ny ch ni n vn lin quan ti dao
ng h thng.
V mt k thut hc, i vi mt h thng, phi ly vic iu khin c lm tin
. V th h thng i vi s bin i ca tn hiu chuyn vo hoc cc loi nhiu c
n nh hay khng v iu kin lm cho n n nh nh th no, l ht sc quan trng.
V s chuyn ng chu k ca h thng (dao ng) cn nghin cu su mt s vn
trng thi khng n nh v kh khn ny. Trng thi khng n nh c hai loi hnh:
mt l trng hp tn hiu ra pht tn n v hn, hai l trng hp dao ng.
nhim cng cng (s tng ln mt chiu ca vt cht nhim) c ngha l h
thng trng thi nguy him pht tn. Vn pht tn l mt vn quan trng v
mt sinh thi hc, y ch yu bn v dao ng. Dao ng c chia ra dao ng t
kch v dao ng cng bc, y ch bn v dao ng t kch. Dao ng t kch, ni
bng ngn ng ca sinh thi tc l vn ng chu k ni sinh.
Dao ng ca h thng tuyn tnh: Dao ng m chng ta nghin cu, ni chung l
khng c dao ng tuyn tnh. Nhng kin thc v dao ng tuyn tnh li rt quan
trng i vi vic nghin cu vn ng khng tuyn tnh.
Trong h thng biu din bng phng trnh vi phn hng nht mt bin s khng
dao ng. H thng hai bin s tr ln (hng 2 tr ln trong phng trnh i thnh 1
bin s) cng tc l ni n phi l mt h thng th mi c iu kin tin ca dao ng.
Di y bn v iu kin dao ng ca h thng nh nht (m hnh 2 thnh phn).
dx
dt
= a
1
x + b
1
y + m
1
dy
dt
= a
2
x + b
2
y + m
2

(27)
Vit gn h phng trnh (27) theo y thnh:
) m a m a ( y ) b a b a (
dt
dy
) b a (
dt
y d
2 1 1 2 1 2 2 1 2 1
2
2
= + + + (28)
t nghim ca phng trnh c trng ca phng trnh (28) l r
1
v r
2
th dng
tch phn l:

1 2 2 1
2 2 1 2 rit
2
rit
1
b a b a
m a m a
e C e C y

+ + = (29)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

161
Nu nghim r
1
v r
2
gm c phn o (trng hp nghim l s phc) th pht sinh
dao ng.
Cng thc le (Euler) dng hm s lng gic nh ngha hm s lu tha s o.
e
jt
= cost + jsint (30)
Trong : 1 j =
Nghim r
1
v r
2
ca phng trnh c trng ca phng trnh (28) c tnh bng
cng thc tm nghim ca phng trnh bc 2, ta c:
( )
2
) b a b a ( 4 b a ) b a (
r , r
1 2 2 1
2
2 1 2 1
2 1
+ +
= (31)
t phn trong du cn l D (bit thc) th:
D = (a
1
+ b
2
)
2
- 4(a
1
b
2
- a
2
b
1
) = (a
1
- b
2
)
2
+ 4a
2
b
1
(32)
D < 0 l iu kin ca dao ng, lm cho D < 0, theo cng thc (32) cho thy,
th a
2
v b
1
phi khc du nhau (nu a
2
> 0 th b
1
< 0; a
2
< 0 th b
1
> 0) l iu kin
cn.
Dao ng ca h thng khng tuyn tnh: x l thng nht i vi dao ng
khng tuyn tnh, li khng lm mt tnh cht ch ca n, cn phi c kin thc ton
hc tng i cao.
Di y gii thiu phng php phn tch mt phng pha (mt tng v) cng
khng coi l rt cht ch. y l phng php da vo s phn tch qu o mt phng
pha gn im cn bng phn on xem c kh nng dao ng hay khng v l loi
dao ng no. Gi l phng php phn tch mt phng pha ngha l em nghim ca
phng trnh vi phn mt bin s hng 2 (33) ban u t ln mt phng x - y nghin
cu (thi gian khng biu hin, l phng trnh hin).
x
2
+ ax + bx = 0 (33)
Do , khng thch hp vi h thng hng 2 tr ln. Nu trong phng trnh (33)
m x = y th c th chuyn phng trnh vi phn mt bin s hng 2 v h phng trnh
vi phn hng 2 bin s hng 1, m hnh 2 bin s (hng 1) mi s dng c phng
php ni trn.
By gi, h thng hp thnh bi hai thnh phn x v y, c dng vi phn l:
dx
dt
= P (x, y)
dy
dt
= Q (x, y)

(34)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

P(x, y) v Q(x, y) khng c quan h g vi tuyn tnh v khng tuyn tnh, trn mt
x - y t to ca im 0
dt
dy
, 0
dt
dx
= = l (x
0
, y
0
), im gi l im k d (im cn
bng). Tm x
0
, y
0
bng phng trnh sau y:
P (x, y) = 0
Q (x, y) = 0


(35)
Nh vy mc din t h thng khng tuyn tnh ni, t im (x, y) gn
im (x
0
, y
0
) v cch nhau khng xa, thay (x - x
0
), (y - y
0
) vo x, y ca phng trnh
(34), thng qua khai trin Taylor, b cc s hng bc 2 tr ln i, tc l tm c
phng trnh xp x tuyn tnh nh sau:
dx
dt
= a
1
(x - x
0
) + b
1
(y - y
0
)
dy
dt
= a
2
(x - x
0
) + b
2
(y - y
0
)

(36)
Trong :
0 1
x x
x
P
a =

=
0 1
y y
y
P
b =

=
0 2
x x
x
P
a =

=
0 2
y y
y
P
b =

=
(37)
Thut ton tip theo hon ton ging vi trng hp dao ng tuyn tnh nu trn.
Kh y ca phng trnh (36) l tm ra phng trnh vi phn hng 2 ca (x - x
0
) nh sau:
DP
2
(x - x ) - (a + b ) D (x - x ) + (a b - a b ) = 0 (38)
0 1 2 0 1 2 2 1
Phng trnh c trng ca h thng ny l:
r
2
- (a
1
+ b
2
)r + (a
1
b
2
- a
2
b
1
) = 0 (39)
t nghim ca phng trnh (39) l r
1
v r
2
th

2
4 ) ( ) (
1 2
2
2 1 2 1
2 1
b a b a b a
r r
+ + +
= (40)
162
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

163
Cc s hng quyt nh tnh cht ca nghim trong cng thc trn l:
M = a
1
+ b
2

D = (a
1
- b
2
)
2
+ 4a
2
b
1


(41)
Cn c vo cc du hiu ca M v D, tm ra qu o (x, y) gn im k d, c
trnh by nh hnh 11.5 (ghi ch: phn bit D > 0 khng phi quyt nh du ca M
m phi cn c vo u D M ).
x x x
x
x x
y y y
y y y
M=0 M>0 M<0
D<0
D>0
Ni r thm l, trong vic tnh ton
bng phng php trn, cn ch cc
im sau y:
1. Trng hp D = 0, nu khng xt
n c cc s hng bc cao ca chui
Taylor th c sai s.
2. Trong h thng khng tuyn tnh
khng phi ch c mt s im k d, v
phng trnh (35) khng phi ch c mt
nghim.
Hnh 11.5. Quan h gia du ca biu
thc (D,M) trong m hnh 2 bin s (x,y)
vi dng qu o mt phng pha x-y
gn im k d (im cn bng)
ca nghim gii
3. Trng hp D < 0, h thng thnh
trng thi dao ng qu o ca mt x - y
thnh dng xoy. Cch quay ca qu o
dng xoy ny ph thuc vo du ca b
1
.
Nu b
1
> 0 th quay thun chiu kim ng
h, b
1
< 0 th quay theo chiu ngc li
(ly x lm trc honh).
1) M = a
1
+ b
2;
D = (a
1
- b
2
) + 4a
2
b
1
2) Phn on dng qu o khi nghim thc
D>0 ch da vo du ca M th khng
chnh xc, phi cn c vo du ca nghim
3. M hnh ho my tnh
Th no gi l m hnh ho
M hnh ho l s dng ci khng ging vi i tng nghin cu (m hnh)
nghin cu din bin v c tnh ca h thng c nghin cu, t ny ng ngha vi
thc nghim m hnh.
V loi cng c m m hnh s dng v ngha dng chng tin hnh thc
nghim m hnh ni tit th nht ca chng ny. Trong s nhng loi cng c m
hnh th tnh nng ca my tnh l tt nht, nht l nhng nm gn y, s pht trin
nhy vt ca my tnh in t, dng my tnh tin hnh thc nghim m hnh tr
thnh ph bin, do c th ni ngy nay l thi i ca m hnh ho tc l s dng
my tnh in t tin hnh thc nghim m hnh.
Mc ch ca m hnh ho: M hnh ho i th c hai mc ch (cng dng)
chnh: mt l gim nh gi thit khoa hc, hai l thc nghim thay cho vt thc. Mc
ch u ch yu l cng dng v mt nghin cu. Rt nhiu m hnh lm ra trong qu
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

trnh nghin cu c th khng hon thin, c nhng im khng r rng b chn vi
trong nhng gi thuyt khoa hc ca cc nh nghin cu. Dng my tnh i vi m
hnh c nhng gi thit khoa hc ny, i chiu kt qu thc nghim vi nhng hin
tng bit h thng thc t, c th phn on c tnh hp l v tnh chnh xc
ca nhng gi thuyt y. Khng nhng th, cn c th da vo s quan st kt qu i
chiu, sa li gi thuyt hay gi cho nhng ti nghin cu mi khi so snh vi
thc t.
V thc nghim thay cho vt thc, ch yu l nhng vn v mt ng dng,
y cn ly m hnh ng tin cy lm tin . C khi chng ta ngh n mt thng tin
ng dng ca h thng no , nhng v l do kinh t, k thut, an ton hay mi trng
m khng th tin hnh thc nghim trn h thng vt thc c. Th d nghin cu
phng php phng tr nm bnh o n la, tm hiu phng thc lan truyn trong
phm vi ln ca loi nm ny, trong thc t khng cho php th nghim loi nm ny
bt k vng rung nc no. Hoc khi ly cy trng c hnh thi c th cha c trong
thc t lm mc tiu to ging, khng th dng vt thc thc nghim hiu sut quang
hp; khi nghin cu cc phn ng khc nhau ca h sinh thi cc iu kin kh hu
nhn to, tuy v mt k thut c th lm mc no , nhng cn phi chi ph rt ln
v tn nhiu thi gian, trn thc t l cha th lm c. i vi nhng vn nh th
(nu c c m hnh ng dng ph hp), pht huy tc dng nht vn l thc nghim
m hnh ho bng my tnh.
Phng php ton hc v m hnh my tnh
Phng php x l ton hc (gii tch), theo ngha rng cng l mt loi m hnh
ho trn giy, y ni r thm mt bc phng php ton hc sn c, ti sao cn cn
dng my tnh tin hnh m hnh ho.
Lnh vc cn tnh bng s: H thng c nghin cu thng qua cc biu thc ton
hc, thng tin c x l bng gii tch ton hc c tnh ph bin nht v m bo
chnh xc. Nhng phm vi nhng vn tm c nghim bng x l gii tch li rt
hp, hu nh khng lng trc c. Phi c y cc iu kin cht ch sau y
mi dng c phng php gii tch tm nghim:
1. Phng trnh dng tuyn tnh
2. Phi l t bc 4 tr xung (trong vng 4 bin s).
Nu khng tho mn c 2 iu kin ny, th ch c th da vo cc phng
php tnh ton tr s, ngoi ra khng cn bin php no khc.
D l m hnh ho tho mn c iu kin nh vy, nu l bc 3, bc 4, th
nghim tm c cng ht sc phc tp, nh th m c tip tc tin hnh gii tch
tm nghim th rt tn cng sc.
Tm nghim bng gii tch ni y tc l trng hp thng s (a
1
, a
2
, ...a
n
) ca
h thng no hon ton khng biu th bng s, m l bng ch.
164
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

Gii phng trnh vn c ca h thng, mt bin s y
1
no biu th dng hin
ca cc thng s h thng v chuyn vo.
y
1
= f
1
(a
1
, a
2
.... a
n
, t) (42)
Nu dng hm s v phi ca phng trnh ny gin n, th c th thu c mt
s thng tin v quy lut tng qut, th d: iu kin cho y
i
tr thnh cc i l a
1
=
0,5 a
2
2
= 0, a
3
tng ln ln th y
i
gim theo hm s lu tha... Nhng nu khi dng hm
s v phi rt phc tp, cho a
1
tng ln, cha bit chc y
i
tng hay gim, trng hp ny
cn a cc tr s c th a
1
, a
2
, a
3
,... an vo tnh ton, cng tc l phi lp i lp li
nhiu ln tr s mi lm r c xu th ca hiu qu a
i
. y, tm nghim bng gii
tch cng rt kh khn. Ni tm li, tm nghim ca h thng tng i phc tp, v th
p dng cch tnh ton tr s l con ng tt nht, ngoi ra khng c cch no khc.
Tnh bng s v my tnh in t: Nhng phng php tnh ton tr s c quan h
mt thit vi vic m hnh ho l phng php Hone (Horner), phng php Graep
(Graeffe), phng php Bestu (Bairstow) gii phng trnh bc cao; i vi h phng
trnh nhiu bin s th c phng php kh, phng php Gao-Sy en (Gauss - Seidel),
phng php ng t; i vi phng trnh vi phn thng, c phng php Runge -
Kutta, v.v... Nhng phng php ny v mt l lun u c th tnh bng tay. ng
thi, mi vic tnh ton tr s phn nhiu cn nhng th tc tng i phc tp, h
thng cng phc tp th lng tnh ton cng ln. Nht l vi thc nghim m hnh ho,
vn d c c im nhiu loi hnh ton trong m hnh, thng km theo mt khi
lng tnh ton lp i lp li ht sc ln. Tnh ton bng tay tt nhin cng c, nhng
trn thc t ht sc chm. Ngy nay, a s cc nh nghin cu u c lin h vi trung
tm tnh ton no , h hon ton khng phi t tay mnh tnh ton tr s, ht thy u
nh my tnh in t.
My tnh ch s v my tnh m hnh ho: My tnh in t c coi l cng c tt
nht cho vic m hnh ho, c th chia ra lm hai loi: my tnh ch s (Digital) v my
tnh m hnh ho (Analog). Trong thc nghim m hnh ho h sinh thi nn chn loi
no, hin nay vn cha xc nh. ng v gc ca kiu m hnh ho, so snh tnh
nng ca my tnh, sau ny s ni n, nhng vic chn my tnh, ngoi vic xt tnh
nng ca n ra, mu cht vn l vn loi my tnh no dng thun tin hn. Nu ch
nhn vo s lng my tnh, th kiu m hnh ho khng khc vi kiu ch s, nhng n
thin v b mn k thut hc v phn nhiu do c nhn s dng.
My tnh kiu ch s c s ngi s dng tng ln rt nhanh. y ch yu ni v
my tnh kiu m hnh ho.
Cu trc v cch s dng my tnh kiu m hnh ho:
My tnh kiu m hnh ho c pht trin t thit k my gii tch phng trnh vi
phn. M hnh ng ca h thng m chng ta nghin cu c din t bng phng
165
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

trnh vi phn, cho nn c th d kin l vic s dng my tnh kiu m hnh ho trong lnh
vc nghin cu sinh thi s ngy cng tng. y ch nu nhng iu ti thiu v my
tnh kiu m hnh ho.
u im ca my tnh kiu m hnh ho
- B phn ca h thng thc t v m hnh c s i ng r rng: My tnh kiu m
hnh ho p dng phng thc tnh ton song song ng thi, lng tnh ton l mt
lng lin tc nh in th. V th c tnh tng t cao gia c nng ca bn thn my
tnh v c nng ca hin tng t nhin. Th d, s quang hp ca l, cc hot ng
sinh l nh s chuyn vn sn phm quang hp n cc c quan khc, s ht m ca
b r, trong h thng thc t c tin hnh song song ng thi. Nhng my tnh kiu
ch s v mt c nng khng xt n tnh ng thi ca cc hin tng t nhin, m
tnh ton ln lt tun t theo mt th t nht nh. Xt v im ny, my tnh kiu m
hnh ho ging vi h thng thc t (hin tng t nhin), tin hnh tnh ton song song
ng thi. Do , trong nhm b tnh ca my tnh kiu m hnh ho, c th ch ra mt
cch r rng ci no i ng vi c quan quang hp, ci no i ng vi s ht m ca
r v ng h hiu th h s trong my tnh i ng vi cc thng s. c im ny
gip rt nhiu cho vic tm hiu m hnh my tnh.
- Tnh ton nhanh: my tnh kiu m hnh ho vi phng thc tnh ton song song
ng thi, cho nn d l h 2 phng trnh vi phn hay h 100 phng trnh vi phn th
thi gian tnh ton cng khng khc nhau, u trong khong 30 giy l tm ra gii p.
V th cng l h thng ln th tc tnh ton tng i cng nhanh hn so vi my
tnh kiu ch s.
Trong h sinh thi, phn nhiu l thng s cha bit, cho nn khng th khng tnh
th lp i lp li nhiu ln. Nh vy, tc tnh ton nhanh l mt u im.
- Thao tc n gin: Nu ch l thao tc tnh ton, th ch cn 10 pht l nh
c, r rng l n gin d hc. Trong qu trnh tnh ton, va xem xu th ca p n,
khng cn ngt qung s tnh ton m vn c th thay i trnh t ca n hoc tr s
thng s, thao tc ny cng rt n gin. Nh vy, ht sc c li cho vic tin hnh thc
nghim m hnh ho nhm kim nh gi thuyt khoa hc. Ngoi ra, v l tnh ton in
th, vic ghi li v quan st p n cng tng i d dng, gi tin my khng qu t,
cng l mt u im.
Nhc im ca my tnh kiu m hnh ho:
- chnh xc thp: sai s ca b tnh tuyn tnh l 0,1 - 0,01%, b tnh khng
tuyn tnh c hi cao hn, l 0,2 - 0,05%. Trng hp do nhiu b tnh t hp li, tho
lun mt cch khi qut sai s tng hp ca chng khng n gin. Cng vi vic m
rng h thng, s b tnh phi s dng tng ln tng ng, cn d tnh n l sai s tt
nhin cng tng ln.
- V thi gian tnh ton ca my tnh, xt n sai s ca n, cn khng ch trong
phm vi thch ng (loi tc thp l 10s - 1 pht), qu di hn hoc qu ngn hn
phm vi ny, sai s u tng ln. Do , nu khi thi gian ca h thng thc t vt qu
166
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

167
hoc b xa phm vi ny, th cn bin i thi im ca n (lm cho h s ln ln hoc
nh i mt cch tng i), lm cho thi gian ca my tnh trong phm vi thch hp
th mi tm c p n. Nhng mt h thng no , chng hn nh h thng mt ci
l, trong cng mt h thng, c hin tng kch thch ca phn t sc t quang hp cp
10
-10
s, cng c hin tng ho gi ca l cp thi gian my tun l, nu m hnh ho
cng mt m hnh, th sai s ca n l mt vn ln. H sinh thi c c trng l cng
tn ti nhiu thnh phn hp thnh c nh s thi gian khc nhau.
- Lng tn tr (b nh) nh: Nng lc ny thp hn nhiu so vi my tnh kiu
ch s. V th, khi dng s liu o lng chiu sng mt tri, nhit khng kh lm s
liu u vo, cn thm ph kin chuyn dng bn ngoi my tnh (th d nh b c
ng cong chng hn).
- t hm s c bit: thnh phn c sn dng c log [x] sinx, cosx, x
2
, x .
Ngoi ra, c mt s hm s phi dng b pht sinh hm s tu kiu ng gy, th
dng lc no lm lc y. Thao tc cho d kin vo tng i phin phc.
- Nng lc phn on logic km: im ny gn y c ci tin, nhng vn khng
so c vi my tnh kiu ch s.
Trin vng ca my tnh kiu m hnh ho
Cho n nay, my kiu m hnh ho vn cn tn ti nhng nhc im k trn.
Nhng gn y, do s tin trin v thit b cng, khc phc c mt s nhc im,
pht huy c mt tt vn c ca my tnh kiu m hnh ho, v ngy cng hon thin.
Ni tm li c hai hng ci tin:
1. My tnh kiu m hnh ho tnh ton t ng;
2. My kiu hn hp.
My tnh kiu m hnh ho tnh ton t ng: Trong m hnh khng tuyn tnh, nu
mun gii vn tr s ti u,
vn bin tr v tr s cho,
th ngoi cch lp i lp li nhiu
ln th sai ra, khng cn cch
no khc. Loi kiu my tnh ny
c thit k da vo mun c
gng a vic th sai nh vy
tr thnh t ng ho. Hin nay,
hu ht cc my tnh u c kh
nng tnh ton t ng.
M
Th rng
Mo rng
0
25
50
75
100
125
150
1855 1860 1870 1880 1890 1900 1910 1920 1930
Hnh 12.5. Bin i chu k ca s c th
Bin i s c th thng k sn bn th rng
v mo rng Canada
Chu k khong 10 nm (Mac Lulich, 1937)
Theo ti liu ca Ito v Kritami (1971)
y tnh kiu hn hp: Kt
hp my tnh kiu m hnh ho
v my tnh kiu ch s c nh
vi nhau, pht huy cc mt tt
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

168
in nay ngi ta u dng loi my tnh kiu m hnh ho m vic chng trnh
ho
4. Phn tch h thng mt s m hnh sinh thi
vi my tnh tin hnh phn tch
h th
ng ca m hnh lin to thc n (m hnh tnh chu k):
k (hnh 12.5).
Ngu
n to thc n 1 (m hnh Lotka Voltera, hnh 13.5):
hc

ca chng - tnh tc cao ca kiu m hnh ho, tnh x l s liu v tc chnh xc
cao ca kiu ch s, nng cao tnh nng ca c my tnh.
H
c th tin hnh t ng.
Hin nay, ngi ta dng phng php ton hc
ng i vi m hnh c th. Ch yu l thc tp phng php phn tch, khng t
vn ngha sinh thi hc ca kt qu phn tch.










Phn tch
C nhng ng vt d sinh, s c th ca chng bin i c tnh chu
yn nhn gy nn hot ng chu k nh vy c: thuyt kh hu (mt thuyt v dao
ng cng bc lin h vi tnh chu k ca hot ng ca mt tri), thuyt bnh cy,
thuyt stress (tc l tm nguyn nhn t bn trong qun th ca mt loi no , thuyt
quan h lin to thc n gia cc loi (m hnh Lotka Voltera, hnh 13.5) ... y ch
yu cp n thuyt Lotka - Voltera, tho lun v h thng lin to thc n v hin
tng chu k.
M hnh li
Cc nh ton hc Lotka (1925) v Voltera (1926) ln lt a ra m hnh ton
v s bin ng s lng c th ca loi.
xy k x k
dt
dx
2 1
=
(43)

y k xy k
dt
dx
3 2
=
(44)
x y
(+)
k
1
x k
2
xy
k
3
y
t mi ng vt i ng v bt m
k
1
x
1
k
2
y
0
k
1
y 10
x
x

+
-
+
-
-
+
+
- -
B phn ghi B phn
(x)
1) M hnh hnh v
o
Hnh 13.5. M hnh lin to thc n (m hnh Lotka - Voltera)
3) S s dng my tnh
kim m hnh ho
n hc 2) M hnh t
hi
(y)
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

169
Trong : x - S c th (hoc khi lng) ca loi lm mi
y th ca loi bt m i x.
x do b bt n hay t l
ng trnh (43) v k
2
ca
ng du khc nhau, x v y y l khi

- S c
k
1
- T l tng t nhin ca x, k
2
l t l gim ca
tng ca y, k l t l cht ca y (k
2
ca ph
3
phng trnh (44) biu th b
lng, nn cng du).
(43) v (44) l h phng trnh vi phn khng tuyn tnh, cch gii gii tch chnh
xc rt kh khn. Nhng n l m hnh hai yu t, cho nn c th dng phng php
gii tch mt pha ni trn.
Trong phng trnh (43) v (44) t 0
dt
dy
0
dt
dx
= = v th im k d (y
0
, x
0
) ca
h thng ny l:
k
3

x =
0
k
2

k
1
k


y
0
=
2
(45)
t v phi ca phng trnh (43) l P, v phi ca phng trnh (44) l Q, lm
thnh xp x tuyn tnh g ca cc phng trnh, cc h s ca phng trnh
c trng nh sau:
k c M v D nh sau:
M = 0 + 0 = 0
(47)
0
: h th ng ny gn t h
thng h vng trn
khi d o tm p s th c kt qu nh hnh 13.5.
n im k d


P
x
= k
1
- k
2
y x = x
0
, y = y
0
, a
1
= 0
P

y
= - k
2
x x = x
0
, y = y
0
, b
1
= - k
3




C n c t qu ca cng thc (46), bit th

D = (0 + 0)
2
+ 4 k
1
(-k
3
)
= - 4 k
1
k
3
<
Phn on th t hnh 16.5 c th th y im k d, l m
m dng xoy nhm ng tm. M hnh ny t sc d dao ng, c i
ng my tnh kiu m hnh h
Hnh 14.5 l gii p tm c tri qua x v y thi gian khi t k
1
= 0,3 ; k
2
= 0,6 ;
k
3
= 0,3. Tr s tuyt i ca cc thng s thiu cn c ca bt k loi hnh sinh thi hc
no, ng nhin khng c coi trng lm.
Q
x
= k
2
y x = x
0
, y = y
0
, a
2
= k
1
= k
2
x - k
3
x = x
0
, y = y
0
, b
2
= 0

(46)
Q
y
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

170

Hnh 14.5. Bin ng chu k ca ng vt lm mi (x) v ng vt bt mi (y)
tm c theo m hnh Lotka - Voltera (munekata, nguyn hnh),
(t l thi gian phn tch my tnh kiu m hnh ho l 1 giy)

Gi thi h sinh thi
i nm tnh ton. Trn hnh
,4 nm). Nh trn ni M = 0,
cho
nh nh trn phi ch gn im k d (x
0
, y
0
) mi c c tc
dng
Ghi ch: x
0
= 3, y
0
= 3, k
1
= 0,3, k
2
= 0,6, k
3
= 0,3
t 1 giy ca thi gian my tnh tng ng vi 1 nm ca
thc t, th k
3
= 0,3 c ngha l t l cht ca y ly 30% m
ny cn c th c c chu k l khong 4,4 giy (4
nn c th ni h thng ny l mt dao ng lin tc, khng suy gim cng khng
pht trin. Nhng trn hnh c th thy n c hi suy gim. l sai s sinh ra do vic
xp x tuyn tnh gn im k d. Trong trng hp ny, im k d x
0
(= k
3
/k
2
) = 5 ; y
0

(= k
1
/k
2
) = 5, cho nn x v y u tim cn vi 5 khi t . Hnh 14.5 gn vi ng
cong ca hm s sin.
So snh hnh 14.5 v hnh 15.5, nu lm cho k
2
nh i, th thy: 1) thi gian chu k
ln ln; 2) dao ng tr thnh kiu pht tn. Do h thng ny l khng tuyn tnh, th
tc lm xp x tuyn t
. Nu tm thi khng xt n im ny, th M phn on tnh n nh ca dao ng
v chu k dao ng nh sau:
M = (k
1
- k
3
) + k
2
(x - y) (48)
Chu k
2 1
k k
2
= (49)
T cho thy, nu k
3
nh i, th M > 0, ngha l d pht tn hay thi gian chu k
ln l nh u ln thc nghim m kin
ca dao ng lin tc (M = 0) i th l:
k
1
- 2k
3
= 0 (50)
ln. Ngoi ra, chnh l s iu i y tnh cho thy, iu
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

171

Hnh 15.5. Lm nh t l cht (k3) ca ng vt bt mi (y) th thi gian
s c th thp ko di ra, xut hin pht sinh ln dng mch xung (pulse)
Ghi ch: 1) x
0
= 5, y
0
= 3, k
1
= 0,3, k
2
= 0,6, k
3
= 0,13
2) x m rng gp 2 ln
Hnh 16.5 l m
3
nh
i, kho h i,
bin p ni
trn ng ca loi lm mi x v loi n
mi y nh


Hnh 16.5. Gii p my tnh kiu m hnh ho m hnh Lotka - Voltera khi bin i
t l cht (k
3
) ca y. Nu t l cht nh i, chiu rng ca mt chu k ln ln
Ghi ch: x
0
= 0, y
0
= 3, k
1
= 0,25, k 0, k
3
= 0,10 0,18
t th gii p khi k
3
phm vi 0,10-0,18 (cp 0,2). Khi k
ng cch chu k ln ln, dao ng hng v kiu pht tn. Ngoi ra, k
3
n
dao ng ca x cng thp xung nh cng thc (45). C ba trng h
u i gian, th hnh ly trc honh lm trc th
sau:
x = f
1
(t)
y = f
2
(t)

(51)


0,18
1 giy
7
6
8
von
0,16 0,14 0,12
k
3
= 0,10

5

4

3

2

1

0

2
= 0,5
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

172
Ln lt coi x, y l chuyn vo a

trn
i l ng
thng s, c trng hp quay trn
mt
x,
iu
chn t,
xem
im
xut pht
vo my v th x - y, c th biu th 2
gii p coi l qu tch ca 1 im
mt phng x - y. g nghim t
v.
Hnh 17.5 l nghim tng v ca x,
y; hnh ny cho thy do dao ng suy
gim m tr thnh ng cong xoy
hng tm (im k d) theo s t hp
ca
ng cong khp kn, c khi xoy c
m rng ra ngoi (dao ng pht tn).
Chiu xoy ca h thng ny theo chiu
ngc chiu kim ng h, c th thy
trc c cn c vo b
1
= - k
3
(< 0).
Hnh 18.5 l nhng dao ng u
tin khi bin i k
2
cc tr s. hnh
ny t k = k
3
, nh cng thc (45) cho
thy, khi k
2
bin i, ton b im k d
(x
0
, y
0
) u nm trn ng thng y =
im k d
0) khi k
tin gn v im gc (0,
2
ln ln. Ngoi ra, khi cho
k
2
ln ln, dao ng ta nh nh i,
trn thc t l v im k d tin
gn v im ban u (3, 3), khng
phi l hiu qu trc tip ca k.
M hnh lin to thc n II:
Chng ta tin hnh m hnh
ho hiu chnh (thay i) cc thng
s ca m hnh Lotka - Voltera.
Di y l trng hp th h
h bn thn kt cu k thu
n nh hng nh th no n
vic tm p s. Chng ta tin hnh
nhiu hiu chnh i vi m hnh
Lotka - Voltera, nhng nhng thc
nghim hiu chnh khng phi l
d bn v hiu chnh, m ch l vic
th gii p vn .
0 1 2 3 4 5 6 7 8 9 10

8
7
Hnh 17.5. Gii p bng my tnh loi lm
m t mi y(t) ca m hnh
Lotka - Voltera, dng my tnh v th
Ghi ch:
1 2 3



x
y 10
9

6
5
4
3
2
1
i x(t) v loi b

x - y v trn mt phng
1) k = 0,373; k = 0,687; k = 0,367
t t = 0; x = 2,89; y = 3,00
2) Vng trn suy gim cng gn im k
d (5.35; 5.42) cng tim cn, d v
hnh nn khng tnh ht
0 1 2 3 4 5 6 7 8 9 10
10
H

nh 18.5. th dao ng u tin v trn


m t x-y khi thay i cc h s bt n (k
2
) trong
hnh Lotka - Voltera. im ban u ca x
v y cch xa im k d (k
3
/k
2
, k
1
/k
2
) th
dao ng u tin cng ln
9
8
7
6
1
2
3
4
5
m
Ghi ch: khi k1, k = 0,3; k = 0,55 1,00,
2 3
t = 0, x = 3,0 y = 3,0
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

173
t hn ch
bt k
ca loi, tc l khi tc ly thc n ca loi sinh vt lm mi x c hin
tn
c hiu chnh c dng sau y (tham kho hnh 17.5, 18.5):
Lng ly thc n (tc ) ca mt loi sinh vt no , khi lng sinh vt t, gn
nh thnh t l vi s lng ca n. Nu lng sinh vt rt nhiu, mt nhn
no c tc dng, th ni chung, lng ly s t n im nh. Th d quan
h gia tc quang hp (tc ly CO
2
) v din tch l (= lng thc vt) ng l
nh vy.
M hnh Lotka - Voltera t c hai loi u c tc ly thc n thnh t l vi
s lng
g t n im nh ni trn, quan st ton b hnh ng ca h thng s bin
i nh th no.
Gi thit hin tng n nh ny c th biu th bng hm s hipe - bn, th m
hnh ton hc
) P ( x k xy k
bx a
x
k
dt
dx
4 2 1

+
=
(52)
) Q ( y k xy k
dt
dx
3 2
=
(53)
S hng k
4
x trong phng trnh (52) l tc cht ca x (ni chnh xc ra cn gm
c tc h hp v bi tit). Trong m hnh Lotka - Voltera, trong k
1
ca phng trnh
(43)
k
2

(54)
bao hm c k
4
, cn phng trnh (52) li lm cho tc ly thc n tr thnh
phi tuyn, do cn thit phi tch ra s hng k
4
x. Theo cch lm trc, trc ht l
lm xp x tuyn tnh gn im k d, tin hnh x l ton hc, c im k d (im
n nh) ca h thng ny l:
k
3

x
0
=

k
1
k
2
- k
2
k
4
a - k
3
k
4
b
y
0
=
(k a + k b) k
2

(55)
2 3
Quan h chuyn vo ca x ch p lm cho hm s biu th cn
bng ca y tr nn ht sc phc t o y
0
bng k
1
/k
2
trong m hnh ni trc,
cng
3
2 3
)P
2
(< 0) (56)
hi phc t
p. Cn c the
im
tc l ch xem xt y
0
c quan h vi k
1
, im ny c th d kin c.
Bit thc ca dao ng nh sau:
- k k k b
1 2
M =
(k a + kb
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

174

2
) b

+
+
=
2 1 4 3
1
3 2
k k k ) b k (
k (
k
k a k (
3 k
D (57)
M < 0 cho nn c th phn on tt c u l h thng n nh suy gim. Cn c
cng thc (57) phn on D m hay dng cng kh phin phc. N i,
gii tch h thng tng i phc tp.

cng thc (57) cho thy: a, b, k
3
,
k
4
l n
ng ca n cng tr nn kh khn hn.
Tron
nh). V
k
4
=
+
2
a k
+
2
3 2
) b k a
2
3
2
2
2
b k k
h trn n










T
n ln th D d g, cng tc l dao
g trng hp ny, dng thc nghim
m hnh ho s nhanh hn nhiu. M thc
my tnh kiu m hnh ho ca h thng
ny nh trong hnh 19.5. S dng b phn
ca b chia, tng ng vi b phn ca
hm s hipebn vi x chuyn vo.
Hnh 20.5 l hiu qu bin i h s b
i vi h thng (khi b ln ln th hm s
hipebn t rt nhanh n im
0, trng hp b = 0 ca hnh ny hon
ton ging vi kt cu ca m hnh Lotka -
Voltera trnh by phn trc. T hnh
ny cho thy: khi b ln ln, th dao ng
tt rt nhanh. Nh vy l nht tr vi kt
qu ca gii tch (b ln, th D > 0). Ni
x y
(+)
k
2
xy k
3
y
ng vt

ng vt
bt mi
1) M hnh
h h
3) S s dng my tnh kim m hnh ho
k
1
2) M h h ton h n c

= y k xy k
dt
2 1

=
=
dx
x k xy k
x k
dt
dx
4 4 2
1
bx a
Hnh 19.5. M hnh lin to t n II
-
hc
x
0
k
2
y
0
k
1
y 10
x
(x)
B phn
(y)
k
0
+
b
a
b = 0,0
10 giy
1 von
Hnh 20.5. Hiu qu s b (h s
lm tng cng khng tuy ca
hm s tng sin i) i vi dao
b = 0,1
b
ca h
n tnh
h vt m
ng ca m hnh lin to thc n x (t)
b = 0,2
b = 0,3
= 1,0
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

175
ng ln
hn,
y ly tun hon m n gin ho lm th d, ni v nh hng ca h s
tc
c bng tay, lm
ton b ng dy thnh tuyn tnh, (1), (2), (3), ca hnh ny, trong h thng thc t
bng ngn ng ca sinh thi hc, tc l do hiu qu ca bn thn, trong h thng cu
thnh ly loi tng nhanh ngay n im nh lm loi sinh vt mi, th khng d g
pht sinh hin tng chu k. Hnh 21.5 l hiu qu ca t l cht ca sinh vt mi x. Do
t b = 0, nn ging vi m hnh Lotka - Voltera. Nhng khi hiu qu ca t l tng v
t l cht ca x l s m, v im ny th khc nhau. Khi t l cht ca x tng cao ln,
tc suy gim ca chu k s t rt nhanh n im cn bng, ng thi lng y
trng thi n nh cng gim theo. T l cht ca x hu nh khng c nh hng n tr
s n nh ca x. ng nhin l ni trng hp t l cht khng phi l hm s ca
thi gian. Khi t l cht ca sinh vt mi t n mt gii hn no tr ln (cng tc
l khi hi ln hn t l tng ca sinh vt mi), khi sinh vt mi v sinh vt bt n u
tr thnh 0, h thng ny s mt.
M hnh tun hon vt cht (vn tr s n nh ca m hnh tuyn tnh)
H thng tun hon vt cht so vi h thng lin to thc n, l mt h th
cho nn ng dy ca n cng phc tp hn, y l mt i tng rt tt v gii
tch h thng. Tuy nhin, s hiu ca h thng thc t, ni v tng b mn, c
tch lu kh, nhng nhn v ton b th c th li rt khng y .

im xut pht
0,25
0.1
0,10
0,16
k
4
= 0,0
0 2 3 4 5
ht (k
4
)
7
6
5
4
2
1
3
x
Lng sinh
y











vt lm mi
1 6
Hnh 21.5. Gii p pha khi thay i t l c
ca x trong m hnh lin to thc n II

(thng s h thng) i vi trng thi n nh ca h thng.


Hnh 22.5 l hnh v ca m hnh cn tho lun. tnh ton
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

c
2 3
(58)

Khi tr s ban t on trng thi
qu , ri cui cng s t n trng thi n nh (hnh 22.5).
Ch cn gii phng trnh tuyn tnh nh th l c. Nghim tm c nh sau:
k
coi l khng tuyn tnh. Chuyn m hnh v dng ton hc, tr thnh h phng
trnh vi phn 4 bin s bc 1 sau y:
x
1
= a - k
1
x
1
+ k
5
x
4
- k
6
x
1
- k
7
x
1
x = k x + k x - k x
1 1 2 2 2
x
3
= k
3
x
2
- k
4
x
3
x
4
= k
2
x
2
+ k x - k
5
x
4
+ k
6
x
1 4 3
Cy trng
lu nm
ng vt
n c









u v tr s n nh khc nhau, sau khi tri qua m

Nu ch tho lun tr s n nh, th trong phng trnh (58) :


x
1
= 0 , x
2
= 0, x
3
= 0 , x
4
= 0
a
x
1
=
7


a k
1
x
2
=
k
7
(k
2
+ k
3
)

X
2
X
3
m
X
1

Cht h
trong t
u c
X
4
3
2
1
3 2
k
3
x
4
k
1
k
4
x
3 k
1
x
3
k x
k
2
x
1
3
x
a
k
Hnh 22.5. M hnh m gi thit.
Trong 1), (2), (3) vn phi c kt cu
tun hon
ng (
mi lin h ngc (khng tuyn tnh), y
tm khng xt, ton b lm cho chng gn
vi tuyn tnh; a, k
1
- k
7
l h s (thng s),
x
1
- x
4
l bin s ph thuc v thi gian
2
x
1
176
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

177
1
k
3
x
k
7
+ k
3
)k
4
a k
3
=
(k
2

x
4
=
k
7

k
5
a k
1
+ k
6
Gi thit m hnh ny l im sau y c xc lp:
1/ Ch cn khng thay i l ng chuyn vo chuyn ra ca h thng, hng s
tc t
ch
ng con ca h sinh thi ng rung, m hnh sinh trng
ca im m hnh ho. Phn ny trnh by trong
Gio
ng, th thng th
i
ng cc

(k
1
, k
2
, ... k
6
) trong ng vng d thay i nh th no, lng m trong
u ni chuyn vo chuyn ra cng khng thay i.
2/ D c nng cao t sut tc ng vt n cy trng, lng gi N ca ng vt
vn c hn nht nh.
M hnh sinh trng ca cy trng:
Cy trng, coi l h th
n c rt nhiu thit k v thc ngh
trnh Sinh thi Nng nghip.
0 20 40 60 80 giy
Thi gian tnh ton
Hnh 23.5. g my tnh
kiu m hnh ho theo m hnh tun hon m
t hu c trong t
Cy
x
1
x
2
x
3
x
4

8
7
6
5
4
2
Ch
Von
ng vt n c
trng lu nm
m trong t
3
Th d kt qu tnh ton bn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

TM TT
Thng thng cc bin php chia nh s gip chng ta n gin ha h thng.
Tuy nhin kiu suy lun tnh cht ca phn t nh suy ra c im ca ton h
thng i khi khng cho chng ta kt qu mong mun. Phng php tip cn
h thng theo quan im sinh thi hc s cho chng ta mt cch nhn tng th
hn v qun th cy trng v h sinh thi ng rung. M hnh ha c s
dng nh mt b cng c hu ch nghin cu h thng. ng thi n cn c
tc dng bo trc cho chng ta bit trc cc tc ng ca h thng. Trong
thc t cc nh khoa hc thng s dng m hnh ha cc mi quan h gia
cy trng, dinh dng t, c di, nh sng nghin cu chc nng v cu
trc ca h sinh thi ng rung. thc hin m hnh ha, ngi ta biu din
cc mi quan h di dng i s hoc h thng cc phng trnh ton hc
dng tuyn tnh hoc phi tuyn tnh. Hin nay, ngi ta dng phng php ton
hc vi my tnh tin hnh phn tch h thng i vi m hnh c th. Chng
ta c th tham kho thm v d c th v m hnh sinh trng ca cy trng
trong gio trnh Sinh thi hc Nng nghip.

CU HI N TP
Dnh cho sinh vin i hc v sau i hc:
1. Hy trnh by u nhc im ca bin php chia nh?
2. Ti sao ni sinh thi hc mang tnh tng hp?
3. Hiu th no l h thng? S khc bit gia h thng c hc v sinh hc?
4. Trong qu trnh phn tch h thng phi lu nhng im g?
5. Mc ch ca nghin cu h thng l g?
6. Hy trnh by cc c trng c bn ca m hnh h thng VAC?
Dnh cho sinh vin sau i hc:
1. Th no l trng thi n nh?
2. Th no l h thng tuyn tnh?
3. Lm th no x l dao ng ca h thng khng tuyn tnh?
4. Th no gi l m hnh ha my tnh?
5. Hy nu u nhc im ca m hnh ha my tnh?
178
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
GII THCH THUT NG
1. Sut phn x (Albedo): T l ca thng lng nh sng nhp x v lng
dng nh sng phn x. Trong qun th cy trng sinh trng pht trin tt, t
l ny c tr s bng khong 0,2. Tr s ny thay i nhiu theo gc nhp x.
2. T l nc dng ln u (Primary water use percentage): Ch tiu dng
quan st mc s dng lng nc ma ln u ca rng, ng rung, t
cn... Tr s ny khng tnh nc dng li.
3. Hiu sut chuyn i nng lng (Energy conversion efficiency): T sut
m quang hp c nh c trong s nng lng nh sng m thc vt hp
th. Qun th la nc pht trin tt th hiu sut chuyn i nng lng l
khong 7%.
4. Tr khng khuch tn (Diffusion resistance): Cng dng in gia hai
im t l thun vi hiu in th, t l nghch vi in tr. Suy ra, lng vt
cht chuyn vn (khuch tn) gia hai im t l thun vi hiu nng vt
cht gia hai im t l nghch vi tr khng khuch tn vt cht . Tr
khng ny gi l tr khng khuch tn, tng ng vi s nghch o ca
tc trao i.
5. Mi trng (Environment): nh ngha khi qut l iu kin ngoi cnh ca
sinh vt, tc l tng ho nhng iu kin bn ngoi c quan h vi i sng
sinh vt.
6. Trng thi n nh nht (Climax): S hp thnh qun x trong thc b thin
nhin din bin theo thi gian, cui cng t n loi hnh qun x n nh
nht trong iu kin chi phi ca ni - ch yu l iu kin kh hu. Loi
hnh qun x nh th gi l qun x cao nh (n nh nht). Qun x cao
nh s tip tc tn ti vng lu di nu khng b ngi, ni la hay ho
hon ph hoi.
7. Qun x (community): V mt sinh thi hc, thut ng ny biu th th cng
ng ca nhiu loi sinh vt, i tng thc vt th l qun x thc vt (Plant
community), i tng ng vt th l qun x ng vt (Animal community),
c khi gm c thc vt v ng vt th gi l qun x sinh vt. Ngoi ra, cn
c vo i tng nghin cu, gi l qun x c di, qun x cy trng...
8. Lng hin cn (standing crop): Lng sinh vt (biomass) tn ti mt din
tch no trong mt thi gian no , gi l lng hin cn, thng thng
biu th bng khi lng vt cht kh.
1
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
9. Thng lng hin nhit v tim nhit (Sensible and latent heat flux): Trong
s nhit nng a vo mt mt no , nhit lu ng do s chnh lch nhit
gia mt v khng kh chung quanh n gi l hin nhit (lng dng),
nhit lu ng vi hnh thc tim nhit trong bc hi gi l tim nhit (lng
dng).
10. Tc trao i (exchange velocity): Mt loi vt cht no trong mt
dung mi no , khi chuyn dich (trao i) t im A n im B, lng
chuyn dch trong n v thi gian, n v din tch mt ct thnh t l vi
hiu nng ca loi vt cht ti im A v im B, v h s t l c th
nguyn ca tc (cm/s), cho nn gi l tc trao i.
11. Thnh phn hp thnh (component): H sinh thi ng rung do qun x
sinh vt trong ng rung v nhng yu t khng phi sinh vt nh nh sng,
CO
2
, nc, t, thnh phn dinh dng v c... cu thnh. H thng c cu
trc c hc v chc nng nh th gi l h sinh thi ng rung.
12. Qun x sinh vt ng rung (field biome, biotic, community in field): Tn gi
tng qut cc thnh phn sinh vt nh cy trng, c di, su hi, vi sinh vt, ng
vt... sinh tn trong h sinh thi ng rung. Khng bao gm cc thnh phn
khng phi sinh vt.
13. Qun th (population): Cu thnh bi nhng c th sinh vt cng loi.
14. Cu trc hnh hc tng tn (geometrical structure of a canopy): S phn b
khng gian ca cc b phn trn mt t ca cy trng. Mc din tch b
mt cc c quan c biu th bng hm s phn b thng ng v hm s
biu th phng php tuyn dng trn b mt cc c quan.
15. H thng (system): nh ngha ca h thng l 1/ t hp ca nhiu thnh
phn hp thnh c quan h qua li phc tp; 2/ l mt chnh th (th thng
nht) trn ngha no .
16. M hnh ho (simulation): Tuy nhng yu t (thnh phn) s dng hon ton
khc vi h thng i tng nghin cu hin thc (h thng thc), m hnh l
s n gin ho h thng thc nhng phi c cc c tnh quan trng nht ca
h thng thc, ngha l m hnh c ch to ra c cu trc l lun ph hp
vi cc yu t ca h thng thc.
17. Sinh vt d dng (heterotrophic organisms): Ch nhng sinh vt ly dinh
dng vi hnh thc hp cht hu c, da trc tip hay gin tip vo sinh vt
t dng. Nhm sinh vt ny li c chia ra: sinh vt dinh dng hon ton
ng vt, dinh dng hoi sinh, dinh dng k sinh...
18. Sn lng thun (net production): Lng cht hu c tch tr li c th
thc vt; cng tc l lng quang hp tr i lng h hp.
2
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
19. Sut ng ho thun (net assimilation rate - NAR): Thc o hiu sut
quang hp ca thc vt tc l tc sinh trng ca thc vt, biu th bng
trng lng vt cht kh mi n v din tch l trong mi n v thi gian.
20. Bc x thun (net radiation), cn gi l cn bng bc x: L tng ca hiu
gia bc x ngn (bc x ngoi) chiu trn mt mt no v phn x ca
n.
21. Cu trc sn xut (productive structure): S phn b theo phng thc ng
ca cc c quan trn mt t ca qun th thc vt (cy trng); c th chia ra:
cu trc khng gian ca tp on l (sn xut vt cht kh) v cu trc khng
gian thn cy - c quan khng ng ho.
22. Nng sut (productivity): Sn xut vn d l thut ng ca kinh t hc, trong
nng nghip l ch s thu hoch m ngi ta thu c trn t ai. Cho nn
sn lng thu hoch c trn din tch nht nh gi l nng sut. Tng ng
nh vy, tng lng ton b cht hu c m mt nhm sinh vt sinh ra, gi l
nng sut sinh hc.
23. H sinh thi (ecosystem): H thng ng thi do nhng sinh vt cng sng
trong mt vng no v nhng yu t mi trng khng phi l sinh vt
(nh sng, nc, t...) trong vng to thnh. Tu theo i tng v vng
nghin cu c gi l h sinh thi rng, h sinh thi dng sng, h sinh thi
ng rung...
24. Tng khng kh gn mt t (Air layer near the ground): Phm vi chu nh
hng s ma st gia khng kh v mt t gi l tng gii hn khng kh.
di tng ny, phm vi 40 - 50 cm trn mt t gi l tng khng kh gn mt
t.
25. H thng tuyn tnh v khng tuyn tnh (linear and nonlinear system):
Phng trnh dng din t h thng gm nhiu s hng, bin s v o hm;
trong cc s hng u l bc nht th gi l phng trnh tuyn tnh (phng
trnh i s tuyn tnh hay phng trnh vi phn tuyn tnh), ngoi ra th u l
phng trnh khng tuyn tnh.
Th d:
1/ 0 = a
1
y
1
+ a
2
y
2
+ a
3
y
3
2/
3 3 2 2 1 1
y a y a y a
dt
dy
+ + =
3/
t 3
e y 5
dt
dy
t
2 dt
y 2 d

= + +
l phng trnh tuyn tnh;
3
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
1/
3 3 2 2
1 1
1
1
y a y log a
y b 1
y
a 0 + +
+
=
2/
3 3
y 1 b
2 1 1
y a e a y a
dt
dy
+ =


3/ t sin y a
dt
dy
2 dt
y 2 d
1
2
= + +


l phng trnh khng tuyn tnh
c im ton hc ca phng trnh khng tuyn tnh l khng th xc lp
"nguyn l trng hp", do khng th gii bng ton hc tuyn tnh (tm
nghim bng gii tch h phng trnh, phng php c in ca phng trnh
vi phn, phng php bin i Laplace, phng php bin i Fourier); tr
trng hp c bit ra, ni chung l khng th gii tch tm nghim ca h
thng khng tuyn tnh.
26. Tng sn lng (gross production) hay sn lng th: Ton b cht hu c
m qun x thc vt thng qua quang hp sn xut ra gi l tng sn lng,
cng tc l tng ca sn lng thun v h hp.
27. Hng s mt tri (solar constant): Cng bc x mt tri khi n tng
gii hn khng kh gi l hng s mt tri, c tr s l 1,94 cal/cm
2
/pht.
Hng s mt tri bin ng trong phm vi 1% tr xung.
28. Nhit truyn dn trong t (soil heat flux): Nhit nng t mt t xm nhp
vo trong t gi l nhit truyn dn trong t. Thng c nh gi bng
phng php phn tch phn nhit, phng php tm truyn nhit, phng
php truyn dn nhit.
29. Sinh vt t dng (autotrophic organisms): Nhng sinh vt c th t tng
hp c cht hu c cn thit cho c th cht v c; chia ra: sinh vt t
dng hon ton (thc vt quang hp) v sinh vt t dng v c ho nng
hp. i biu cho loi trc l thc vt mu xanh lc; loi sau l nhng vi
khun khng th trc tip li dng nng lng mt tri, nhng thu c nng
lng thng qua oxy ho cht v c.
30. Cn bng nhit lng (heat balance): Tng ton b nng lng (nhit) ra v
vo trn mt mt no trong thi gian nht nh, theo nh lut bo ton
nng lng th bng 0. S ra vo ca nhit gi l cn bng nhit lng.
Bc x thun trn mt t, lu ng hin nhit, lu ng tim nhit, truyn
dn nhit trong t l ni dung ch yu ca s cn bng nhit lng v c
gi l cc yu t cn bng nhit lng.
4
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
31. Hm s phn b l (leaf distribution function): Hm s biu th cu trc
hnh hc ca tp on l, cng tc l hm s biu th phng php tuyn dng
trn mt l. Phng ca php tuyn do gc thin nh v gc phng v quyt
nh. Thc t l s phn b tn din tch l tnh ton theo gc thin nh v
gc phng v ca nhng l khc nhau.
32. Nn nng nghip t ngp (flood farming): Do l lt ca cc sng ln, khi
lng ln bn t, cht lng ng c chuyn xung h lu, hnh thnh
vng t ai mu m ngp nc. Nn nng nghip pht trin y gi l
nng nghip t ngp, l mt hnh thi ca nn nng nghip nguyn thu t
thp.
33. Vi khun quang hp (photosynthetic bacteria): Vi khun lu hunh (sulphur
bacteria) mu v mu lc xanh li dng nng lng nh sng ly cc cht
hu c i loi nh axit bo cp thp hay H, cc cht sunfua lm th cung cp
H sinh sng, cng thc phn ng l: 2H
2
S + CO
2
2S + CH
2
O +H
2
O.
Nm 1940, Gest v Kamen t trong Rhodospirillum rubrum pht hin loi
vi khun ny c kh nng c nh m. Vi khun quang hp c nh m kh
nhiu trong iu kin k kh v c chiu sng.
34. H hp nh sng (light respiration): Thc vt C
3
c h thng h hp nh
sng nh CO
2
trong iu kin chiu sng; thc vt C
3
h thng nh vy
cng nhiu hn. Trong iu kin nng O
2
tng i thp, h hp nh sng
b c ch. Thc vt C
3
khi nng O
2
h thp th quang hp ca n tng ln.
35. Sn xut vt cht (sn xut vt cht kh) (dry matter production): Thc vt
nh hot ng quang hp ca b phn mu xanh lc sinh ra cht hu c
sinh trng. S to ra cht hu c ny gi l s sn xut vt cht. Thng
thng biu th bng trng lng kh ca c th thc vt, cho nn cn gi l
sn xut vt cht kh.
36. Thng lng (flux): Mt loi vt cht no (hoc mt loi nng lng no
) khi vn ng (lu ng) trong mi trng, lng chuyn vn ca vt cht
trong n v thi gian, n v din tch trn mt mt mt no th gi l
thng lng.
37. T s Bowen (Bowens ratio): Nhit a vo mt mt no , trong c
phn nhit lu ng dng hin nhit (H) v phn nhit lu ng dng tim
nhit (IE), t s H/IE ly tn ngi xng ra n, do c tn l t s
Bowen ().
38. Cn bng nc (water balance): Cng nh cn bng nhit lng, tng s ra
v vo ca nc th rn, th lng, th hi trn mt mt no trong mt thi
gian nht nh no bng 0. Ni c th l s ra vo ca nc: nc ma,
nc chy trn b mt t, trao i nc gia mt t v tng di, bc hi
nc b mt, gi l s cn bng nc.
5
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
39. Nn nng nghip nng ry (slash and burn farming): Nn nng nghip
nguyn thu, t rng hoc ng c xong, ngi ta trng cy (ch yu la
nng, sn, ng) t mt n vi nm. Lp tro trn mt t l ngun cung cp
dinh dng v c cho cy trng. Nng ry s c b ha sau khi t mt
mu v ngi dn li pht mt mnh nng mi.
40. Nc hu hiu (available water): Lng nc m thc vt c th ht c
t trong t, tng ng vi lng d t lng gi nc ln nht (h s
ho) m t c th gi c. Hm lng nc hu hiu khc nhau tu loi
t, t tht v t tht pha st cao hn t ct v t st.
41. Lng nc cn (Water requirement): tm hiu nc cn dng trong thi
gian sinh trng ca thc vt, ly tng lng nc ht chia cho tng khi
lng vt cht kh th c lng nc cn gi l h s bc hi.
42. Ch s din tch l (leaf area index, LAI): T s gia ton b din tch l trn
mt t nht nh v din tch ca mt t .
43. Bao b mch c dip lc (chlorophyllons bundle sheath): Trong bao b
mch l ca thc vt C
4
nh ng, la min... c cht dip lc, gi l bao b
mch c dip lc. Thc vt C
3
nh la nc, m, u tng... khng c loi
m ny.
44. H s khuch tn dng xoy (turbulent transfer coefficien): S lu ng ca
th lng nu vt qu mt iu kin no th tr thnh lu ng c nhiu
ch xoy, gi l dng xoy. Trong loi lu ng ny, s khuch tn ca
vt cht no thnh t l vi dc nng v h s khuch tn ca vt
cht . H s khuch tn trong dng xoy gi l h s khuch tn dng xoy.
6
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
MC LC
Trang
Li ni u 3
Chng I
NG RUNG V SINH THI HC NG RUNG 5
1. Khi nim chung v sinh thi hc ng rung 6
2. Qu trnh hnh thnh v pht trin ng rung 8
Tm tt 12
Cu hi n tp 12
Ti liu c thm 12
Chng II
CU TRC V CHC NNG CA H SINH THI NG RUNG 13
1. Cn bng lng nhit v cn bng nc ca ng rung 14
2. Mi trng t 22
3. Mi trng sinh vt 24
4. Cu trc ca qun th cy trng 26
5. Cu trc mi trng ca h sinh thi ng rung 35
6. Quang hp ca qun th cy trng 48
7. S sinh trng ca qun th cy trng 57
8. S cnh tranh ca h sinh thi ng rung 61
9. Nng sut ca h sinh thi ng rung 65
10. M hnh ho h thng h sinh thi ng rung 69
Tm tt 82
Cu hi n tp 82
Ti liu c thm 83
Chng III
S VN NG CA H SINH THI NG RUNG 85
1. Din bin ca ng rung 86
2. S bin i hnh thc sn xut nng nghip v ngha sinh thi ca n 90
3. Trng cy thch hp vi vng t trng 100
4. S cn bng nng lng ca qun th cy trng 104
5. S chuyn ho nng lng ca ng rung 108
6. Tun hon vt cht ca ng rung 110
Tm tt 125
Cu hi n tp 125
Ti liu c thm 126
1
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Chng IV
IU KHIN H SINH THI NG RUNG 127
1. ngha v phng php iu khin h sinh thi ng rung 128
2. iu khin qu trnh ca h sinh thi ng rung 130
Chng V
K THUT HC H THNG CA H SINH THI NG RUNG 139
1. Sinh thi hc v k thut hc h thng 139
2. Chun b ton hc m t v phn tch h sinh thi 148
3. M hnh ho my tnh 163
4. Phn tch h thng mt s m hnh sinh thi 168
Tm tt 178
Cu hi n tp 178
Ti liu c thm 179
GII THCH THUT NG 180
TI LIU THAM KHO 185


















2
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn












nh xut bn nng nghip
167/6 - ng Phng Mai - ng a - H Ni
T: 5763470 - 8521940 FAX: (04) 5760748
chi nhnh nxb nng nghip
58 Nguyn Bnh Khim - Qun 1 - Tp. H Ch Minh
T: 8297157 - 8299521 FAX: (08) 9101036





















3
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn







Chu trch nhim xut bn
NGUYN CAO DOANH
Bin tp v sa bn in
NGUYN TH HI
Trnh by, ba
TRN V











63 - 630
NN - 2006
- 202/78 - 06



In 1.000 bn, kh 19 x 27cm ti Xng in NXB Nng nghip. Giy xc nhn ng k
KHXB s 360-2006/CXB/202 - 78/NN do CXB cp ngy 9/5/2006. In xong v np lu
chiu qu III/2006.
4
Tai lieu chia se tai: wWw.SinhHoc.edu.vn
Tai lieu chia se tai: wWw.SinhHoc.edu.vn

You might also like