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THERMOTROPISM OF ROOTS
CONTRIBUTIONS FROM THE HULL BOTANICAL LABORATORY 192
SOPHIA ECKERSON
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
INVESTIGATION
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
on permeability
The effectof temperature is shownin the
accompanyingfigures.The ordinatesgivethepercentageweight-
1914] ECKERSON-THERMOTROPISM OF ROOTS 259
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
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
0.42- --?
o Ii
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
10 20
FIG. 5.-Phaseolus primaryroots; the sucroseand glucose curvesare
multiftorus:
from
identical400 to250 C.
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.
+ 0 + -
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.