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 THERMOTROPISM OF ROOTS
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 192
SOPHIA ECKERSON

(WITH SIX FIGURES)

I. The thermotropic ofcertainrootsat temperatures

curvatures
from I10 to 400 C.
oftheserootscausedbychange
II. The changesin permeability
in temperature from10 to 400 C.

Introduction
Our knowledgeof thermotropism is in a somewhatconfused
state. We know littleabout the thermotropic responseof stems.
A fewscatteredobservations have beenmade,indicating thatstem
tipsturntowardthesourceofheat at low temperatures and away
fromit at hightemperatures.Moreexactinvestigations havebeen
madeof the thermotropic curvaturesof but
roots, herethere seems
to be no uniformity of behavior. Underthe influence of unequal
temperature on twosides,therootsofdifferent speciesreactdiffer-
ently. Some rootsgive positivecurvaturesat low temperatures
and negativeat high temperatures;othersshow only positive
curvatures,and stillothersgiveonlynegativecurvatures. It was
withthepurposeof clearingup theseapparentdiscrepancies and
thehopeoffinding thephysicalmechanism of thecurvatures that
thepresentinvestigation was undertaken.

ofroots
curvatures
Thermotropic
LITERATURE

of roots are
The two importantpapers on thermotropism
WORTMANN'S (Io) and KLERCKER'S (2). WORTMANN investi-
gated fourspecies: Ervumlens,Pisum sativum,Zea Mays, and
Phaseolus multiflorus.He concludedthat roots have positive
thermotropism and negativeat hightempera-
at low temperatures
Botanical Gazette, vol. 58J [254
I914] ECKERSON-THERMOTROPISM OF ROOTS 255

tures,that is, above 350 or 400 C. The one exception he found

was Phaseolusmultiflorus, whichgave negativecurvaturesfrom
220 to 500 C., but no curvature below 220. The secondaryroots
of P. multiflorusreactedpositivelyat low and negatively at high
temperatures.WORTMANNthought, therefore,that the primary
rootsmust have a positivethermotropism, althoughhe was not
able to demonstrate it.
AF KLERCKER studiedPisumsativum, Helianthus annuus,Faba
vulgaris,and Sinapis alba. In the firstthreespecieshe observed
only negativethermotropism; whileSinapis alba gave positive
curvatures fromW40to 290 C., and no curvaturesat thehighertem-
peratures. PORODKO(7) studiedthe thermotropic curvaturesof
roots at temperaturesfrom400 to 700 C. He obtained negative
curvaturesand thoughttherefore
that rootshave only negative
thermotropism.'

INVESTIGATION

Method.-Theapparatusused is a modification of GANONG'S

thermostat. A zinc trough,20 incheslong,havinga
differential
zincboxattachedat each end,was heatedat one endby an electric
coiland cooledat theotherendby a freezingmixture. This gives
a fairlyeven gradation from580 to 50 C. The electriccoil has low,
medium, and highadjustment, so thatthetemperature gradientin
thetroughcan be increasedor diminishedas desired. Threesuch
thermostatswerein use at one time.
The seedlingschosenforthisstudywereRaphanussativusand
Pisumsativum. The troughwas filledwithsterilized Sphagnum
This was foundto be-thebestmedium,sincein it the rootsgrow
perfectlystraight. Seeds weresown in the troughat definitely
spacedintervals. Whenthe rootswereI . 5-2 cm. long,the ther-
mostatwas broughtto the desiredtemperatures and the response
I In work described in a very recent article, HOOKER (HOOKER, HENRY D.,

Thermotropism in roots,Plant World 17: I35-I53. QI94) obtainedno reactionin roots

set in I . 25 per centagar. He concluded,therefore, that the curvaturesof rootswhen
grownin sawdust,as in the workof WORTMANNand KLERCKER, are due to hydrot-
ropism. The methodsof experimentationused are subject to criticismand the
conclusionsare erroneous.
2
GANONG,W., Plant physiology,p. 207. i908.
256 BOTANICAL GAZETTE [SEPTEMBER

of therootsobserved. In anotherseriesof experiments,

seedlings
intotheheatedthermostat.
grownat 200 C. wereput directly
Curvatures.-Whenseedlingsof Raphanussativushave been in
thethermostat 2 hours,therootsshowthefollowing reactions:
Positive No curvature Positive Negative
7-I5 C. I6-23 C 24-36' C. 38-5IO C.

At thehighertemperatures thecurvatureoccursveryquickly.
Rootsgrownat 200 C., whenputintothethermostat at 450C., give
a positivecurvature in 5 minutes. This quicklygoes overintoa
negativecurvature,so that within20 minutesthereis a strong
negativecurvature.The explanationof thisdoublecurvature, of
course,is thatas the temperature of therootrisesto 380 C. there
is a positivecurvature;whenthe root temperature has reached
450 C., a negativecurvaturebegins. It shouldbe remembered that
thesideoftheroottowardthehotend ofthethermostat is always
at a slightlyhighertemperature than the oppositeside. At the
lowertemperatures the roots react less rapidly,requiringI. ,-2
hoursforcompletereactionat 7-I5? C.
TABLE I
THERMOTROPIC CURVATURESOF ROOTS

Positive No curvature Positive Negative

i. Raphanus sativus....... 7-I50 C. I6-23 C- 24-360 C 38-5IO C.

2. Pisum sativum. 8-I5 C I7-290 C. .34-50 C.
(WORTMANN) 8 . 5-3IO C . . . 34-50( C.
3. Sinapis alba (KLERCKER) I4-290 C. . .......... None
4. Helianthusannuus
(KLERCKER) .None ... I 5-400 C.
5. Phaseolus multiflorus
primaryroots(WORT-
MANN) . ... ......... . None 8-220 C. ............ 22-50 C.
6. , secondary roots
(WORTMANN)........ IO-?o C.* . . . . ?.-40O C.*

* WORTMANN obtainedpositivecurvatures at 40 C., but madeno

at io? C. and negativecurvatures
at theintermediate
investigations temperatures.

The rootsof Pisum sativumare less sensitiveto temperature

changesthan thoseof Raphanussativus. Afterone hourin the
thermostatall the roots at temperaturesfrom340 to 500 C. showed
negativecurvatures;and in 2.5 hours8o per cent of those at
19141 ECKERSON-THERMOTROPISM OF ROOTS 257

temperatures from80 to I50 C. gave positivecurvatures. At

thetemperatures fromI70 to 290 C. therewereno curvatureseven
after9 hours.
Table I gives the thermotropic curvaturesof five species,
selectedto showthevarious" types" ofreaction. Wherethedata
have been obtainedby otherworkers, thenameis givenin paren-
theses.
In attempting to locatethemechanism of thermotropiccurva-
tures,one mustbear in mindthevariouspossibilities, or "types"
ofreaction,shownin thetable. The rapidityof thereactionindi-
cates thatit is not a growthphenomenon.It is knownthat the
permeability ofprotoplasm increaseswithincreaseoftemperature.
The rangeofpermeability changewas notknown,norwas it known
whetherthereis such greatvariationwith speciesas would be
necessaryto explainthe curvaturesgivenin table I. However,
permeability changesand consequentturgorchangeswere con-
a
sidered possiblefactor;accordingly thepermeabilityoftheroots
at temperatures fromI00 to 400 C. was determined for the five
species.
II. Permeability
EFFECT OF TEMPERATURE ON PERMEABILITY
In 1902 RYSSELBERGHE(8) foundthatthepermeability of the
protoplasm of epidermal cellsof Tradescantia discolor
to dissolved
substances(glycerine,potassiumnitrate,and urea) increaseswith
thetemperature fromo0 to 300 C. In I905 LEPESCHKIN(3) found
thatthevolumeofliquidextruded by hairson theleavesofPhaseo-
increases
lus multiflorus from 00 to 200 C., and decreasesfrom200

to 350 C. Duringthe extrusionthe osmoticpressureof the cell

sap decreases,indicatingan increasedpermeability to solutesas
wellas towater. In i9o8 LEPESCHKIN(4, 5, 6) established thefact
thatpulvinalmovements of leaves are due to theeffectof lighton
the permeability of the protoplasmof the pulvinalcells. Light
increasesthepermeability, withconsequentdecreasedturgorpres-
sure(decreaseofvolume). Darkeningdecreasesthepermeability,
withconsequentincreaseof turgorpressure(increaseof volume).
In i910 TR6NDLE (9) foundthattheleavesofBuxus sempervirens
are morepermeableto sodiumchloridein lightthanin darkness;
 258 BOTANICAL GAZETTE [SEPTEMBER

also witha 32 C.-P. electriclamp as lightsource,he foundan

up to a certainlightintensity
increaseofpermeability (50 cm.from
thelamp),and thena decreaseat thehigherintensities (35-10 cm.
fromthelamp).
METHOD

The methodfordetermining permeability is essentiallythatof

LEPESCHKIN (6). This methodis based on the factthat theper-
meability ofprotoplasm to sucrosedoes not changeundervarying
conditions,whilethepermeability topotassiumnitratedoeschange.
That is, with increasingpermeability, higherconcentrations of
potassiumnitrateare requiredto produceplasmolysis.
Weightmolecularsolutionswereused throughout;thepercent-
ages in any seriesvariedby 0.02 mol. The testsofpermeability
at thetemperatures from20?tO 500 C. werecarriedon in a constant
temperature oven,electricallycontrolled;thosebelow200 C., in an
ice chest. Of course,all werein darkness. Seedsweregerminated
on filterpaper in largePetridishes. Whenthe rootswereabout
I.5 cm. long,theywereput in coveredwatchglassescontaining
the solutionsand leftfor20 minutes. Then theywereput on a
slidein a dropofthesolutionand observedas quicklyas possible.
Repeatedexamination ofsectionsof rootsplasmolyzedat vari-
ous temperatures showed that thedegreeofplasmolysis oftheroot
hairsis an exactindication of thedegreeofplasmolysis of thecor-
ticalcellsoftheroot. Whentheprotoplasm drawnback
is slightly
fromthetipoftheroothair,theoutertworowsofcorticalcellsare
slightlyplasmolyzed.When the protoplasmof the youngerroot
hairis beginning to breakup intoseveralparts,thewallsofall the
corticalcells of the corresponding part of the root are slightly
shrunken.In completeplasmolysisthe protoplasmof the root
hairsis roundedup into threeor fourparts,and the corticalcells
are plasmolyzed.The criterion thenforslightplasmolysisof the
rootis thatcondition of the roothairin whichthe protoplasm is
slightlydrawnbackfromthetip.
DATA

on permeability
The effectof temperature is shownin the
accompanyingfigures.The ordinatesgivethepercentageweight-
 1914] ECKERSON-THERMOTROPISM OF ROOTS 259

molecular solution which produces slight plasmolysis. The

increase or decrease of permeabilitywith change of temperatureis
shown by the increase or decrease of these concentrations,the
plasmolyzingconcentrationof sucrose remainingconstant.
The permeabilityof roots of Raphanus sativus (fig. i) increases
fromio1 to i80 C.; does not change fromi80 to 240 C.; increases

0.42 II I- I I -

- III~,al /os
ejle
-.O _ -_7
Suc-rosel7-r- l -- -u
-~l
_ l _
- _
-
ap0H>
-

0.1 _ _ _ _}
lii
j1 _ _ TemOeralure 0 till
E~e3
0 a0 20ape-20 30
40 50
FIG. i.-Rapkanus sativus: curvesshowingthe effectof temperatureon the per-
meabilityof rootsto potassiumnitrateand glucose; ordinatesindicatethe percentage
wt.-mol.solutionsproducingslightplasmolysis; dots indicate temperaturesat which
determinationsof plasmolysisweremade.

0.42111111

_
S uc rose

0.I8 03

0.10I Tempera ture 4-

.
0 1 0 20
402 _ 0O 50

FIG. 2.-PisUrn sativunn: the potassiumnitrateand sucrosecurves are identical

from 250 to 300 C.

from 240 to 400 C., and then decreases. The turning point in
permeabilityto glucose is at 420 C. The drop in the sucrose curve
above 420 C. suggests a decreased osmotic pressure due to exos-
mosis. To test this, roots were put in distilled water at 450 C.;
after 20 minutes the water gave a sugar test with Fehling's solu-
tion. Thus at 420 C. there is an increased permeability, in all
probability caused by coagulation of the protoplasm.
 260 BOTANICAL GAZETTE [SEPTEMBER

The permeabilityof rootsofPisum sativum(fig.2) to potassium

nitrate increasesfrom60 to I50 C.; does not change fromI5? to

0.42- --?

o Ii

. r 10 O-2 30O ? 4050

FIG. 3.-Sinapis alba: the potassium nitrate and glucose curves are identical
from 400 to 450 C.

0.4

1 IITempcra/ure = ...t

0
. _
42
10 20 30_ 40 50

T
the potassiumnitrateand glucosecurvesare identi-
iF
FIG. 4.-Helianthus annus:
cal from400
0.26|E||||||CI to450 C. o

03_ I 'me'/r I - - I |-l @|||||

10 20
FIG. 5.-Phaseolus primaryroots; the sucroseand glucose curvesare
multiftorus:
from
identical400 to250 C.

236 C.; and decreasesfrom300 to 400 C. The sucrosecurve shows

that the osmotic pressure is lower at 450 C. than at 400 C.
 I9I4] ECKERSON-THERMOTROPISM OF ROOTS 26i

ofSinapis alba (fig.3) increasesfromIO' to 25

The permeability
or 300 C., then remainspracticallyconstantto 450 C.
Helianthusannuus (fig.4) showsno increasein permeability

FTII
withincrease of temperature,but a decrease from200 to 40? C.
0.4; lll
IS ros e- I I

?2t -,
l s i e tX

0..1 I Te;njeratue I 30
1 10 0 30 40 5
FIG. 6.-Phaseolus multiftorus:secondary roots; the potassium nitrate and
sucrose curves are identicalfrom350 to 400 C.

The primaryrootsof Phaseolusmultiflorus (fig. 5) show no

fromI30 to 250 C.; whilein thesecondary
increasein permeability
increasesfrom60 to 250 C. Bothhave
roots(fig.6) thepermeability
a decreasingpermeabilityfrom250 to 350 C.

COMPARISON OF THERMOTROPIC CURVATURES AND PERMEABILITY

CHANGES

To showtherelationbetweenthermotropic curvaturesand per-

meabilitychanges,thedata givenin tableI are givenin tableII
TABLE II
THERMOTROPIC CURVATURES AND PERMEABILITY VARIATIONS OF ROOTS

+ 0 + -

i. Raphanus sativus ..... fCurvatures

.sativus. 7-150 C.
C. I6-23' C. 24-36? C.
C. 24-40? C. 38-5I0
C.
C.
Permeability IO-I4 I8-24e 40500

isu saivu ....Curvatures 8-i50 C. 17-29' C......34-5OO C.

2. 2.Pisum sativum.{Permeability 6-IS C. 15-35 C .35-45 C.

3. Sinapisalba.. f...Curvatures (K)

I4-29O C...........None
{Permeability IO-3O0 C. 30-450 C......None
4. Helianthusannuus. . { Curvatures (K) None ...... C I5-40o C.
iPermeability None 12-260 C...... 20-4O C.
5. Phaseolus multiflorus fCurvatures(W) None 8-22? C. . 22-5 C.
(primaryroots)... . aPermeability None 13-20 C. . 25-40 C.
6. - (secondary fCurvatures(W) io- ?? C.?-4OO C.
roots). Permeability 6-250 C ......... .... 25-40? C.
262 BOTANICAL GAZETTE [SEPTEMBER

togetherwith the temperatures at whichthereis increasingor

decreasingpermeability, foreachspecies. In thetable,+ indicates
positivecurvaturesand increasingpermeability;o indicatesno
curvatures and no changein permeability;- indicatesnegative
curvatures and decreasingpermeability.
The data showthatthepermeability increaseor decreaseparal-
lels almostexactlythepositiveor negativecurvatures.In every
case,at thosetemperatures whichcause positivecurvaturesthere
permeability;wherethereare no curvatures,
is increasing thereis
no changein thepermeability;and at temperatures causingnega-
tivecurvatures thereis decreasing
permeability.The slightdiffer-
encesin temperature at theturning pointare wellwithintherange
ofexperimental error.

Conclusions
WORTMANN'S inabilityto obtainpositivethermotropic curva-
turesin theprimaryrootsofPhaseolusmultiflorus is explainedby
thefactthatthereis no increasein permeability.In thesecondary
roots,however,wherehe foundpositivecurvatures,thereis an
increasingpermeability.KLERCKER obtainedno negativethermo-
tropic curvaturesin Sinapis alba; thereis no decreasingpermea-
bility. Also, KLERCKER obtained only negativecurvaturesin
Helianthusannuus; thereis no increasing permeability,therefore
no positivecurvatures.
The permeability of the cells of the rootto potassiumnitrate
and to glucoseincreasesor decreaseswithincreaseof temperature
accordingto thespecies,and fora givenspeciesaccordingto the
temperature.
Withunequaltemperature on oppositesidesofa root,a curva-
tureis producedonlywhenthecellsoftherootare morepermeable
at one of the temperatures thanat the other. Those cellswhich
are subjectedto a temperature at whichtheyare morepermeable
are
to dissolvedsubstances consequently less turgid. This results
ofthetissuesonthatsideoftherootand a consequent
in a shrinking
mechanicalcurvature.Alwaysthemorepermeable sideoftheroot
becomesconcave.
19141 ECKERSON-THERMOTROPISM OF ROOTS 263

Summary
Thermotropic
i. curvatures ofrootsvarywiththetemperature
and withthespecies.
2. Permeabilityof thecellsof therootto dissolvedsubstances
varieswiththesame factors.
3. In everycase thegreaterpermeability is in theconcaveside
of the root; where the thermotropic reactionof therootchanges,
thepermeability also changes.
4. The parallelismbetweenthepermeability and thermotropic
reactionis exact; turgorchangeproducedby permeability change
offers a mechanicalexplanation of thecurvature.
butbyaffecting
5. Heat doesnotact as a stimulus, permeability
as a directfactorproducingcurvature;hence,thermotropism is
nota tropism, butis a turgormovement.
UNIVERSITY OF CHICAGO

LITERATURE CITED
I. JOST, L., VorlesungenuiberPflanzenphysiologie.Dritte Auflage. Jena.
I9I3.
2. KLERCKER, JOHN AF, tObercalorotropischeErscheinungenbei einigen
Keimwurzeln. Oefvers.Vetensk. Akad. F6rhandl. Stockholm48:765-
790. I89I.
3. LEPESCHKIN, W. W., Zur Kenntnisdes Mechanismusder aktivenWasser-
ausscheidungder Pflanzen. Beih. Bot, Centralbl.I9g:409-452. I905-I906.
4. , tOberdie osmotischenEigenschaftenund den Turgordruckder
der Leguminosen. Ber. Deutsch. Bot. Gesells.26a: 23I-
Blattgelenkzellen
23 7. I 908.
5. - , Zur Kenntnisdes Mechanismusder Variationsbewegungen.Ber
Deutsch. Bot. Gesells. 26a:724-735. I908.
6. , Zur Kenntnisdes Mechanismusdes photonastischen Variations-
bewegungenund der Einwirkungdes Beleuchtungswechsels auf die Plas-
mamembran. Beih. Bot. Centralbl.24':308-356. I908-I909.
7. PORODKO, THEODOR,VergleichendeUntersuchungen uiberdie Tropismen.
II. Mitt. Thermotropismusder Pflanzenwurzeln.Ber. Deutsch. Bot.
Gesells. 30:305-3I3. 19I2.
8. RYSSELBERGHE,FR. VAN, Influence de la temperature sur la permeabilite
du protoplasmevivant pour l'eau et les substancesdissoutes. Recueil
Inst. Bot. Bruxelles5:209-248. I902.
9. TR6NDLE, A., Der Einflussdes Lichtes auf die Permeabilitatder Plasma-
haut. Jahrb.Wiss. Bot. 48:I71-282. I9I0.
IO. WORTMANN, JULIUS,tOberden Thermotropismus der Wurzeln. Bot. Zeit.
43:I93-200, 209-2I6, 225-235. i885.