(SANITIZED)SOVIET PAPER ONCTOPHYSIOLOGICAL AND CYTOECOLOGICAL INVESTIGATIONS OF RESISTANCE OF PLANT CELLS TOWARDS THE ACTION OF HIGH AND LOW TEMPERATURE(SANITIZED)

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• Document Creation Date: December 27, 2016
• Document Release Date: February 14, 2013
• Sequence Number: 1
• Publication Date: November 29, 1963
• Content Type: REPORT

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f t
CYTOPHYSIOLOGICAL AND CYTOECOLOGICAL INVESTIGATIONS
OF HEAT RESISTANCE OF PLANT CELLS TOWARD THE ACTION
OF HIGH AND LOW TEMPERATURE.
V. Ya. Alexandrov
(Review of the works of the laboratory of cytophysiology
and cytoecology of V. L. :omarov Botanical Institute,
Akademia Nauk SSSR).
ABSTRACT
Thermostability of plant cells is due to the resistance
of their proteins to denatration, resistance to injurious
metabolic changes, reparatory ca-aacity, and capacity to harden.
Hardiness includes the stability of several :actions and
increases the resistance to several injdriouL. - It
yaries with the tissue and stage ofr 7rowth. The thermostability.
of the proteins is constant in cr? -,plants but changes with
temperature in algae. 21,oLt hardning increases resistance to
several ? injurious factors including heat. The denaturation
theory of injury satisfactorily explains some of the data.
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CYTOPHYSIOLOGICAL AND CYTOECOLOGICAL INVESTIGAT/ONS
OF RESISTANCE,OF PLANT CELLS TOWARDS THE ACTION OF
HIGH AND L(WTEMPERATURE4.
V. Ya. Alexandrov
(Review of the works of the laboratory of cytophysiology and
cytoecology of V. L. Komarov, Botanical Institute, Akademia
NAUK SSSR.)
I. IICRODUCTION
The principal purpose of the laboratory of cytophysiology
and cytoecology is the study of the mechanisms which determine
the resistance of plant cells towards the action of various agents
and also the elucidation of the ro10 of Call IresistandeTinzddaptation
of plant organisms to various external factors.
Up to the present time we had directed our attention to
the reaction of the cell on -the-P-ffectzbf high and low temper-
atures. The general program of this laboratory represents the
continuation of the Leningrad Cytophysiological School which
had been headed by D. N. Nasonov, corresponding member of the
AN SSSR.
A e041-1> (As
Alreedy-ipegiuning-with the 30' a, several research workers
ka.A, s d ,.71. cc: ?(
of this school have accumulated a-l-arge--factual-material on the
reaction of the cells - mainly animal - towards the action of
various injurious and stimulating agents. This material had per-
mitted Nasonov and me to advance a& theory of denaturi injuries
_
and stimulation of cells which was presented in two monographs
LEX 41St
(Nasonov and Alexandrov 1940; Nasonov 1955;A Alexandrov 1960).
---
On the basis of cytological and biochemical data we came
to the conclu on_that tbe stimulants of entirely different nature
11;
takswohn some,definite doses ?roduce in the proteins of protoplasm
COMMAI:
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. ovt_
similarchangesofdenaturing type. The first stages of denaturing
which are connected With the., chemical activation of different
groups in a protein molecule can increase the metabolism in the
a..?..A4zAxeolt,
cell and -shorten.the-bitee-of the processes. They can serve a's
.s 0.c.pcc c etrtr.,i/c Ct.
activators of this-or=that:tactiwity---typi-ea-l--tbr the given type of
!Cr" ci0oh,
cells. Very strong actions during which the denaturW changes
4
are considerable and include the whole protoplasm may create in ?
a_
the cell4conditiom which is not congenial with its normal function.
In such cases the result will be. not a stimulation but an injury
to the cell.
When the injury has not gone too far, it usually is rever-
sible and after removal of the acting agent the proteins of the
protoplasm return to their primary conditions. With the further
increase of the dose, an irreversible injury takes place, the
denaturation of the cell proteins becomes final and the cell dies.
The denaturation theory easily explains such non-specific
characters and injury, like lowering the dispersion of protoplasmic
colloids, increase of 'viscosity, increase of affinity towards
dyes, etc. The non-specific character of injury depends first of
all upon the similarity of the signs of denaturing of protein after
the action of various denaturing agents. The second source of
great similarity in the behaviOr'of the cells after different
injuries is the wholeness of the cell as a system. Due to the
interaction of its parts the final result 'of reaction may be very
similar regardless of different parts where the various agents are
applied. However the different types of denaturation are not
identical and a wholeness of the cell system is not absolute.
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This vi].].
will
agents sometimes
`'th-at
explain/the reaction of the cells towards various
have paracular specific features which are com-
bined with general non-specific signs of injury (Alexandrov, 1948).
The principal points of our conception are as follows:
1. The reaction of the protoplasm of different cells towards
the action of injurious agents, which are different in their
Vt SRA tr-61:0C d P-Cpre
physical and chemical nature, is)ofi=ap-ecific.
2. Together with the non-specific characters of injury there
exist specific pecu3erities which are characteristic of the
injurious action of each given agent.
3. The stimulation and the injury are consecutive phases of
0 vf;
the response of the cell towards the injurinI agents.
4. At the bases of both the injury and the stimulation are
denaturing changes of the protein molecules of different
protoplasmic components.
During the cell injury it is necessary to distinguish the primary
Cl,ttovt
denaturkag which is caused by direct application of the acting
01.40Nt
agent itctthe proteins of the protoplasm and the secondary denaturing
when the agent causes metabblit. changes and these changes in turn
lead tO..the-denaturing of cell proteins. Such a secondary denatur-
o-tiox a
-lug takes place for example during decrease of cell respiration
after the action of certain inhibitors of metabOliim... These
data together with some others have led us to presume that the
natural state of the proteins of protoplasm is maintained actively
by the normal course of cell metabolism.
After studying the physiology of animal cells we arrived at
att4vt
the conclusion/ that the denaturing theory helps us to understand
1
the series of phenomena which takes place in the plant cells
? 1See Gunar (1953) about the similarity in irritability
of plant and animal celW
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II. CTTOPHYSIOLOGICAL INVESTIGATIONS OF THE REACTION OF PLANT
CELLS TCWARDS THE ACTION OF HIGH 'TEMPERATURE
Materials and Methods
The principal ,o'bectis-? ,disour investigations were living
cells from the epidermis of leavas of several plant species. We
have avoided usually the sectioning, or the:pulling-off of the
epidermal layer. The occurrence of various changes in the proto-
plasm of the cells which are connected with a mechanical injury are
often mentioned in the literature. Fefdman (1960) hald showh that
certain plants-had respond to a cutting or puIling-'off of the
epidermal layer by increased resistance of the cell to various
actions including heat. Even in the/Lipper epidermis of the scale
tc411
leaf of onion 141Tidh4t
A
asily remove
fthe cells are very often
injured although the injury may be reversible (Alexandrov and
Gruzova 1960). Taking this into consideration we have studied
-4-4e cfate /co-('
in living conditions the epidermal cells of the pieces of-Ieamas
? 1 A
without separating them from the mesoph11. For microscopic study
of small pieces of leaves it is necessary to have very good illu-
mination according to Keller, water or oil immersion objective with
co/
correction screw and infiltration of the tissues with the medium
in which the material was mounted. The infiltration was done by
means of a simplified method, using a syringe (Alexandrov 1954).
The medium as a rule was -the tpp water.
In some of the, plants the epidermal cells are covered with
a complex sculptured cuticle. Folds and furrows and the deposits
of various substances on the cuticle may prevent microscopic study;
Cf 16
and quite oftenl,through this cuticle. impossible) see the
c-
structure of the protoplast. To overcome this obstacle a method
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CONF1
was developed which in many cases gave very good results (Alexandrov
exony.( th.czo
1962B). Before the-studyunder the miCroscope)a piece of leaf which
sucit 41t_ Liciitc.A. Am it!,
contains thethidk-cuticle is removed from the Water and driedby
a.
filter paper. Then the piece is put on a slide in the drop of
cx.:. t 4-r c, Nvc
liquid With tbe4index of-refraction whinha near V) that of the
cuticle, and a cover glass
-re.-Cr-a.cttvr t
of the indexcotrefraction
the help of, phase contrast
is applied. The preliminary measurements
of cuticlein a number of plants,)made with
microscope according to Orossmants method
(1949),showed that the index usually .varies between 1.500 to 1.540.
tv,dice.s
is itder-of-refradtionmare some of the organiclicoils of-
s-14>s-1: vokt s
CiiliconU(for example PPOY.?4PPINZ.011icc1)... T,n217. are not toxic and the
- _
. ?
Aj , 4
pieces of immerged leafN,remain alive dinseveral weeks. The
A
sculptured structure of the cuticle becomes invisible when the oil
is selected correctly (fig. la and b), and the contents of the cell
is clearly seen. The application of ordinary vemeline oil (020 1.481)
;ray'
in many cases gives good results. The substitution of water
m; ?-re...-rract..../e.
(ND20 1.333) by-the media with he-index-of-reflectibn--whictr-ir? near
u!.
t* that of the cuticle has permitted to-include-es=objects epidermis cedis
from leaves of many species of plants which were absolutely unusabreit
for microscopic 0 examination in water.
2. A Comparison of Different Criteria of Cell Resistance to Heat
In order to evaluate the iwat resistance tomard-the-actionl...of
highrIEEpibrature, it was necessary first of all to select ,a criterion
by which it would be possible to judge the degree of injury of the
cells after a certain dose of heat (Alexandrov, 1955). Such criteria
aA; respiration, photosynthesis, exit of substances into the medium,
represent gmleval categories which characterize the tissue as a
whole. Only after application of different microscopicp. methods
CONFIDENTIAE
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r4 tt /'r
of
f
of investigation, itLqpossible'to judge -ilf-mt'r-he---44--ssalive.
A.
She Suppression of plasmolysis and deplasmolysist.the exit of
? . 0 /0 s.s
pigments from the vacules, tbtohange of vital staining, luminescence
A
of thesChloroplasts, and also of the fluorochromes introduced into
StreaTh
the cell, the, depression of tbe.protoplasmic motion, changes in
er,4 4.1?:1 a-e"
the cytoplasm and the nucleus duringthe phase :contrast atidatua-
dark field microscopy, change in viscosity which is measured by the
;:shap'el.of-pla7smoly-sior brodiafr-iffigaitorr- all these are used as
criteria of injury. Before selecting from this list the most
PL?t.
suitable criterion, it was necessary to compare their indicati-orts-
during the different degrees of heat injury. The experiments were
conducted as follows. -E-ither whole leaves or pieces of leaves
were put inti water heated to a desirable temperature,for-r-experiments
-\
av4=Zhemplaced in a thermostat_for_five_minutes. Only the heat
-
temperature was varied. After this=prelbniiiiry treatment,-;obserita-
c. .4: re pnr-Ope?Y'r I OS at,' e'r,". z., .
Its-i-osertnattez-witaL?thisttrat---1.-ndloot. The; was
measured i/NAarburgq-apparatus. Thi.Ohotosynthesis was measured
by the radiometric method (Zalenskii, Semikhatova and Vosnesenskii,
V
1955). Thevital staining was done with neutral red. As a fluoro-
chrome, acridin orange was used. The changes of luminescence of
the fluorochrome and the chlorophyll were observed in a luminescence
I., -re-f-lecc4J, sS:tre ilv1)4
' microscope indirect light. The,matielef-----Vehe,protoplasnytaas
judged by the motion of the spherosomes. The exit of electrolytes
was determined by means of electroconductivity of the medium. Ttte
V
Ifiscosity was measured by the displacement of the nuclei after
centrifugation. To;.obtainr.the-plasmolysisr-the.7solution-of KNO3
phis 7
was use. For vital staining and study of luminescence the material
was infiltrated with the solutions of the dye and the fluorochrome.
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s
The rotette leaves; of Campanula persicifolia L. and the
leaves of Tradescantia fluminensis Vell, were the main objects in ?
these experiments.. Results obtained are illustrated in figures 2 and
3. The numbers indicate the temperature o' the minute* of heating.
. A
The behavior of each indicator of injury is shown by the corresponding
,z et-r-tet:
striping. The beginning of the stripe for each indicator corresponds
to the the temperature of-heating-after which first deviation from the
41,5 t?
norm is observed. The end of-the-striping indicates the temperature
A
which shows the maximum injury. For example, for ttralmotion; the
photosynthesis and the respiration)the beginning of the stripe would
correspond to elowVing of the process and the end of-the=stripe to
? ctsw:io.,
? its final stopping. During the vital stainingof normal cells, the
dye is accumulated in the vacule and the nucleus and cytoplasm remains
colorless. In the injured cee11811 a reverse process takes place.
The dye colors the cytoplasm and the nucleus and there is no concen-
tration of the dye in the vact4es.
1 After heating for 5 minutes the injury to the epidermal cells
of Campanula persicifolia begins to show at the=temperature 560C,
? with, death at 60?C. This is judged by plasmolysis and staining
with neutral red (fig. 2). If=z,Vo ,judge by luminescence of acridine
4
orange, or by respiration of the tissue, it is possible to conAlude
Aitiz
that the injury takes place at a lower temperature. 4gintirely different
QYC/110, 'Y Q' CC ,"JS I C 511-CCCOI
conclusion could be -ma, 1E-the7motion,of cytoplasm ie-T!!!!rled-
/f
as-an-indicator: After heating to 39oC t7ae-protnplasmic=motionslows
Cv, oster 'let?
heat,- istunce,of the protoplasmic proteins.
The above considerations have to be kept in mind when the
sensitivity of different functions is compared within the limits
of the same cell. Fig. 2 shows that in thercells c.i"! ..7(paryilaYM
c sscct-40-A ? oc.v,
-Liman Campanula persicifolia complete secession of taTa-Lo.,Ion after 0-
A /A
5 minute heating ocours_!!_i!4?._To_l!presqcompletely, eaa
fil;Zo7S7y-enthesis heating to 46? is required. Judging by this indicator,
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E6T-or.yArkp
it is possible to conclude that ttftlittii-an-omotiam is less
heat resistant than tiletwOr014141.?, photosynthesis. However, the
0 v ewke.,X
parench cell is gtaable to restore the metica even after
of heating, whereas time photosynthesis is irreversibly suppressed
A
after heating to 48? (Liutova, 1962). If we were to judge, there-
c
fore, gtOut the sensitivityof these two functions several days
after? the action of high temperature, we would lave come to the
Str-ta wo+13
. conclusion that the-m i
otion s more heat resistant a,scompared
withr,tthe photosynthesis.. .
When plant4 and animal cells are compared some interesting
differences: occurin their abilit, to restore the protoplasmic -
.wovirm after the injurious action of heat. Table II shows corre-
zerl?le
plants. Here we see that the temperature
temperature which stops elle protoplasmic
spending data for certain
varies 4 ?- 9%1, between the
Sty cv?oki
4wtiOn aTterLIminute?a heating, and the maximal temperature
cv.tv.,/ cdez
after which the4motivn could still be restore? : The abilityito
restore . which haili been stopped by heating in-animal
.celts-is considerably smaller. After 5 minutes' heating of the
ciliated epithelium of frog, the zone of the reversible suppression
of motion was only 1.2? higher o than the temperature which stops the
.t,Ae
motion of ttzt cilia (41.7o). InAmollusk Unio pictorum and in
es s
Paramecium caudatum this zone was smaller than 10 (Alexandrov, 1955).
The cause of such a difference apparently is as follows:
Experiments illustrated in fig. 2 and 3, show that in the plant
cells the dose of heating required to stop the motion is lower than
that necessary for disturbing the selective permeability of the
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pratoplast and to increase the affinity towards dyes by the
cytoplasm and the nucleus. We have shown that the increase in
absorbing capacity of the protoplasm towards the dyes after tilt
cctiolq
action of various injurious agents is the result of the deraturtag
changes af? the pro,plasMic proteins (Alexandrov and Nasonov 1939,
1943; Nasonov and Alexandrov 1940; Brown 1948 a and b, etc.).
Therefore, the proteins of plant cells which are connected with
0
the function of motion of the protplasm are far more sensitive
towards the denaturing action of heating as compared with the main
Pri
mass A0 of proteins of: the cytoplasm and the nucleus. The pre-
p
liminary investigations carried with the animal cells show that
the temperatures which depress the motion, and those which
increase the affinity oT the pro,Olasm towards the dyes are much
closer. This indicates that in ti a animal cells there is no sub-
stantial rigt between the heat resistance of the contractile
-mar of
proteins andhthe main mass of of the protoplasmic proteins. Con-
.?,
c.e.sal?0)1
sequently, the secession of the protplasmic motion (or the ability
A
of cells to contract) occurs in this animal cells with eu4h doses
tk
that injure the whole cell system more as.-7compared.with the
c-a2ca,ttw.t. proTorf*Shk1C,'
secession ofYinotioriarmrotopleam in tala plant cells. Under such
A c), ? t
?
conditions the.-...repa,ratiorr-of,the cell becomes handicapped because
A
it apparently Is, an expression of the living activity of the cell.
Therefore, in certain cases the renaturing of the denatured
protein may take place immediately after the removal of the
denaturing agent -- homodromict(-7-.---) reversibility, for example,
denaturi4 of tripsin and chimotripsin by heat (unitz and
Northrop, 1935). In other cases, the removal of the denaturing
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agent is not sufficient, and more or less complete restoration of
the original native condition could be achieved only after the
additional treatment of the proteins--heterodromic reversibility
17
(see Neurath, Greenstein, *tnam, Erickson, 1944; Putnam 1956).
The restoration of the protoplasmic motion after heating requires
considerable time. In accordance with the dose used, the period
of recuperation may last up to 18 days. In connection with this
it is logical to assume that the restoration of the motion takes
place either by means of a heterodromic reversibility or by sub-
stitution of the denatured protein molecules by newly synthesized
ones. In both cases it is necessary to assume the necessary
participation of the active met4bolism of the cells. Therefore,
'Lk
it is logical to expect that the success of r,,,,a::ation of a certain
function delinds upon the degree of disturbance of the whole cell
4
system;
oft
5. Th?!. Jeat hardening, or the increase cf resistance of
e...xrast
plant cells after ecndt-ng to high temperatures.
The heat resistance of cells is determined not only by the
eth arm-acre' h 'op
heat-xemistance of the cell proteins and by the ability of cells
to regenerate after the action of high temperatures but also by
the increase of resistance in response to tae IQI;i:4ngizby high
temperaturesias our investigations have shown. The information
tO1
in the literature aboutelcapacity of plants to increase their heat
0
resistance after heat treatment is contradictory and based pritily
on Utoi& cytophysiological methods of investigation (Sapper, 1953;
Biebl. 1950; Laude and ChaAgule, 1953; Coffman 1957).
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An increase ot..-he heat resistance of cells in response to
4IttetoottaitionidtbynteTpoDatuP9v:v?--heat hardening--had been
3
established in our laboratory for, more than AO species of plants
belonging to different families. The following problems were
investigated:
on
Q.
(1) Dependency of he-bs of heat hardening from
a) the length of time of heat hardening
b) the temperature of heat hardening
c) the initial heat resistance of the cells
being hardened
(2) Reversibility of heat hardening
(3) Relationship of the hardened cells towards
other injurious agents.
Fig. 8 shows the dependenc:? of the effects of heat hardening
011
of the cells of Tradescantia fluminensis flicm. the duration of
_
hardening at two temperatures, 36.3 and 33.00. In these experiments
IN,?0-02rsQk
every leaf was cut in two. One half of the leaf was placed in0
water at one of the indicated temperatures, the other, control,
immo remained in water at room temperature. After different periods,
cisstz
indicated on the horizontal-line, the temperature which stops
?th..4* protplasmic motion after 5 minutes was determined in both
A
halves of the leaf. As seen from,,tbz fig. 8, a definite increase ix
_ _ . _ .
,of heat resistance t given temperaturears obtained after 30-60 min,/,
The greatest heat resistance was achieved after 36 hours of hardening.
'V;C 1.. t 41,4Z.c-fQ,, occ y-
A,deorepee-wee-A4bserued_along-the-curve, and with still longer periods
Of hardening, the cells had perished. In the control leaves a certain
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-111111116Sal limo
Increase of resistance is also observed after they are removed
from the plant. This has been observed by us in a number of
plants.
v.. five
The dependency* ofAheat resistance of cells on the
temperature to which the cells had been subjected during a constant
period of heating (16-18 hr) is shown in fig. 9. The heat resist-
ance of leaves kept at 18? served as controls. Within the limits
of 1? to 26? a preliminary treatment with heat does not affect the
01.
elvt
heat resistance of cells. The cells respond with tha increase of
heat resistance only after the action of hither temperatures. The
?
vv?aer
maximum effect of heat hardening (increase of 2.1o) occurs im these
A
conditions at 37.50. Still higher temperatures 0 kill the cells.
4 . iect-
CL-A, COcx, ci 10
Bukharin (1958) obtained increase in the breaking point in thetpurves
Proz cf
0--f----Ptratrei.n=coagulaa.,1-ion_ixtA the protoplasm in Lutescens wheat after
A
keeping the plants at 300; however, the breaking point hard-dropped
considerably if plants were kept at 35?.
After tbe-heat hardening)the resistance of the cells
increases not only towards the 5 min heating, but the whole curve
of the heat resistance of cells is displaced. In some plants after
stope
hardening the Incline of the curve (Q10) changes (fig. 10), whereas
in others it remains the same (fig. 11).
he,C
The fact that the cells do not change their,/(resistance
witItilt
tovslartigraleaturing.the wide range of temperatures but begin to
tr ZS
respond by increasing the resistance when the temperature $z reach.iat
. 4 7
e vs -Co Q a k
the injurious zone mzdek.us,presume that the heat hardening is tales ,Tx,41-,,
reaction of the cells execOly towards the injurious action of the
heating. To verify this supposition a series of experiments was set up.
COMM
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In Fig. 12 Me curve (1) shows the influence of temperature
of- ardenin (duration of hardening is 18 hr) upon the intensity
1-11_ ;,e,o
of photosynthesis et-the leaf of Tradescantia f1uminensis.4 Other
1-KAtc.,re ------------
e on
curves represemrt the dependenc: Et-cm the hare:aning temperature
(2) cnd.of_photosynthesis (3).
of heat resistance of protoplasmic motiori' When these curiles are
tfr=
compared, it becomes evident that with th,1 increase a hardening
t c t lay call tm
temperaturelparalleled wbmthe increase of.:7=trh...catert=srf hardenin4
A
there is a definite suppression of photosynthesis.
xf
The depressive action of hardening temperature has been
A
notice in in the experiments of Kisliuk on young plants of cereal
grains.
increase
After 18 hr of hardening at
of heat resistance of plant
0C
al..ggelowth GiWal-to 13-25%, as compared with the controls. Addi-
tional proof that the heat hardening is the reaction of the cells
towards the injurious action of heating can be obtained from -ner.
experiments which determine the connection between the hardening
36.6o together with the
1-
s, there was definite6inhibition
?
temperature and the general effect of hardening fn plants" which
?Yt.
differ by,utkeir heat resistance. These experiments were carried ()kit
tea
with the epidermal cells of ,sheathes of grasses. From fig. 13
01-64_ r
we see that 1attesz=1404-ert-rrerat:,-.11, to increase the cell
resistance per 1 , the hardening temperature required is: for
A
Dactylis omerata L. --300;'Phragmites communes Trin.--36?;
Fanicum miliacaeum L. and Eleusina indica (L) Gaerth. ?40-41?.
The results, obtained agree completely with the differences in the
heat resistance of these grasses: thus, the temperature which
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stops protoplasmic motion after 5 min heating in unhardened
cells of Dactylis glomerata is 44.0?; in Phra.smites communis
is 46.00; in Panicum miliacaeum is 48.5?, and 'in Eleusine indica
is 49.00. Therefore, the cells which are more heat resistant
a.
in the original state require higher temperature., to obtain the a.
effect,of&rdeniT4for the given period of time,.
If ;4) consider that during the heat hardening the increase
?
1-4
ef resistance is a result of adaptation, then not only the
temperature, but also the duration of hardening is essential.
t2,
However, if to consider this process as a reaction towards the
intensity of the heat injury, in such cases the main significance
should be attributed tow c--: the doe of the heating, but not.
the duration. Lomagin (1961) has studied the possibility of
decreasing the time of hardening with the increase Of the
Aemperature ofrdenin. He has received hardening of theq-pdat
? ,
cells fror lea/ epilievivis of Tradescantia fluminensis. Campanula
persicifolia and Chlorophytum elatum R. Br. after4 1 second
?
action of high temperature. One of his experiments is shown
in Table III. During the short time of hardening, the rise -ef
OCCAPIr V Cvl "ki-41 loqd y
resistance is?taiclaee---vepy-fersti, It can be detected already
afiter..5-10 seconds after secezaitif the 10 seconds of hardening.
A
The facts abovementione4 led us to conclude that the
?A"
resistance of cells during the heat hardening
/ increase of heat
4-
Ls the response to the injurious action of heating.
In the abovementioned experiments the effect of hardening
t-n ,...------,,,
was determined by the increase affresistance tuaarcls'heati, as
p,oltop440e. C...fro,,,,,,01----_ `..?......?,
measured by the rftet-i-ork-of,--the_protoplasm. Howev er _the decrease
... . . . ...... .
?....---------Sbs,q lay- -r-e,soltS 1,,,Z1 re. 0 - olvii= .,- Loa 1-* A 1 C -0'1/c.: 0 T C4vt., joartvi-411
1,4 t
( pt?elt et 4^0/01 ....Da cYptis 9/4, eora 1'4 1 ?9.. , d Le ...c ck --., .tA. t...,Yky$h y if i ,
:,...._
...'
__,.... -------
(Z cof ds 0,1 I c43),
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sensitivity towards heating after heat hardening can be deter-
mined akva using other indicators. As compared with the control,
the hardened cells require higher injurious temperature to suppress
A
bite plasmolysis (Alexandrov and Feledman 1958), for liberation of
-co
anthocyanin from the vacuoles (Lomagin 1961), exit of electolytes
? A
into the medium (Derteva), for the suppression of photosynthesis
tosit
(Liutova 1958) and respiration (Liutova 1962), for the distapbance
-tite
of Alink between chlorophyll and chloroplasts with extrusion of the
former and its absorption by the oil globules (Kiknadze 1960).
Thus, tNe heat hardening ? affectittg entirely different components \
of the cell in regard to the action of high temperatures.
An interesting problem is raised in this connection: does the
heat hardening increase the cell resistance_towards_the heat
atso
or does-iit become more resistant towards other injurious agents?
.A
We did not find n the lit eratureXany indicatioD, whether or not the
increase in heat resistance in plant cells is specific. In the
a,
animal cells it was indicated in a number of papers that adppta-
tion of cells to a certain agent can produce an increase in their
resistance towards other stimuli, which can be entirely different,
both physically and chemically (Daniel, 1909; Neuschloz, 1920;
Haffner and Wind, 1926; Orlova, 1941; Paribok, 1948; Trifonova,
1952; Barbashova and Ginetsinskii, 1956; Polianskii, 1957;
Shliakhter, 1959 and others).
The response of cells from the leaf epidermis of Tradescantia
fluminensis hardened for 18 hr at different temperatures, towards
heating, ethyl alcohol, acetic acid, and ammonia, is shown in
-r-1
fig. 14. With the increase of-tNe,temperatureof hardening; the
cell resistance increases not only towards eie heat but also
/c, y // a In 4 t-ke:
10.0 P?rott.,1,n. cow?rit.)(CLivIenvo. ?
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towards -Mae alcohol and acetic acid. In regard to ammonia the
114,e,resuits
cells did not show any definite increase of resistance. In
4
a vt LYt
Table IV - ti WOMB sho, whtdh increased' triz. resistance
Alt
of the cells towards tfte- hydrostatic pressure. Ilte increase of nt
resistance towards alcohol after heat hardening was found in the
cells from the leaf epidermis of Zostera marina (exper. Fel'dman
and Liutova), in the cells of parenchyma of Campanula persicifolia
/?6?
(Kiknadze, 1960), and in Podophyllum peltatum Cprper. Liutov9.c
Our further investigations 1341" -T? it61:-to determine the
time-duration of the effect of hardening. We have found that
this condition is labile, and during the first 24 hr-, a definite
decrease in resistance of cells towards heat takes place
Y
as well as to some other factors.
(fig. 15),
Thus, we have found, that the plant cells may react to a
pyt
large extent by nonspecific reversible increase of resistance
A
towards the action of a high injurious temperature.
1-f---t-e?eet-ts-liter-tiris--eaot from the point of view of the
at=0-4.
denaturitg theory of injury, the results obtained are not un-
1 "It
expected. In order to explain the increase of resistance of the
cell not only towards tte-heat-Cg, but also towards other injurious
agents, often quite different in t4zir nature, it is sufficient
to realize that at the base of the adaptive increase of heat
is
resistance during the heat hardening is a certain stabilization
,/
-factly skp,v4d n%A flc-yQose
of the native state of the protoplasmic proteinsJ,(Aie-x-a,
111_ stst
e 1.? y
r
C e'll-c 6' ..S? d
, c 4
/(CAAC/ Q.1 CcitC/z. 0; 1-1-cal- Acsy-4eneit
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?
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6. The protective reaction of cells at the moment of heating.
As had been indicated above, the right
the curves of cell heat resistance (fig. 5-7)
3r VL.
of tr/i19 motion:ofiProtoplasm Inz:vin the speed of
Straightrsections.of
express the dependenci
heat denaturrag of
certain proteins wh-ich are connected with the-Eunotion--fef motion.
e FQ`r
In most of the objects this dependenck remains only up to-the
temperatures will-eh stop motion during 20-80 min. At some lower
temperatures this dependencv is q.
t-et
.--a--ged-TtEmTmmtad-s. On the left side of the break erf the curve,'
the protoplasmic mmtion is preserved for a lon,srer period as,eom-
--the-higher temperatures. In fig. 16 is given
a curve of heat resistance of the grass Eleusine indica. If the
period of preservation of the-met-ioR in this grass were to depend
Apon the speed of 'the: heat denaturation of proteins in the zone
cs
.Q-%-(yapoi,cte.t
of lower temperatures, then coninuir.g. the line-vid-ehe?left (dotted
A
section) we would expect*Iithat at a temperature of 45.00
for example, example, the .rnotion of the protoplasm would keve stoppA0,after
StN'Ca
47 min. In reality, the met-ixwycontinues for about 3000 min.
at the same temperature.
2.
What are the causes which Affect the break gf. the curve
in the region of 0' moderate heating? To elucidate this problem,
I have studied the condition of the cells at different periods
xvo
after the beginning of the heattactlen and at a temperature some-
what lower than that which corresponds to tinet-"oint-o-f break uct t-11
the curve. The experiments were conducted in a special heat
chamber which was installed on the
-------- -
?-
e-
cv j be-
tage
microscopi. A
EA,
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piec eE lea from Campanula persicifolia was placed in the chamber,
and the same cells were observed continuously. The result of one
of these experiments is given here. The motion of the spherosomes
became quite slow after 40 min. of heating on the mi9Goscope stage,
a
and the temperature of the object had reached 41.0. 411'11 further
1-y1 -Por
rise .af temperature halken stopped and d,*-7.-,--ng the duration of the
cx,7sriment it was maintained at 41.0-41.5?. (The break -of the
curve of heat resistance in this object is at 42.00, see fig. 6).
the
After 90 min. from the beginning of experiment, the motion had
A
stopped completely. Regard-le-5:8 of continuin-g heatingefter 200
min.hor,seral spherosomes began to move again, some ex-
hibiting a forward motion. The motion continued to be more lively,
tke,
and het] approached normal condition after 6 hr. When the heating
was continued further, a second defate-..9s-ipon- of motion took place,
and, finally the cell krad perished. Exactly similar observations
were made in the cells of Phragmites communis (Alexandrov 1956).
These experiments demonstrate the capacity of the cells to
overcome the injury caused by the high temperature, not only after
c Q. S S ekti Yt.
eeeeevien of heating but also during the heating, provided the Mat-
, C
temperature was not too high. The changes in cell metabolism must
occur te?aehieve a repair during the still conti:nuing action of an
injurious agent. These changes Must either compensate or neutralize
the destructions which are caused by the agent. There are several
investigations which deal,-withC.thistkroblem'(Petinov and Molotkovskii,
1956, 1957, 1960; Molotkovskii 1957), but the essential cause still
remains obscure.
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VW= spolas NI ma
At the the temperatures which are indicated to the right of
the breaking point of the curve of heat resistance, the cells do not
show active resistance tcriards the injurious action. Thus, the
breaking point of the curve should he reggrded as a limit, on one
ect 0 la
side of which the cell resists heat denaturin23 and on the other.side?
it behaves as if it were just be. passive protein system.
The resistance, apparently, consists not only in the regeneration
of the denatured protein but also in the increase of its resistence
during the process of heat hardening of the cell.
When the en',Ase of the cell death is compared in the two
regions on both sides of the breaking point, a substantial differ-
ence can be detected in some cases. In the cells killed at bi-#mer
temperature,?-whizh:-ave higher than the point of resistance, neithar
tho.-haodlike plasmolysis nor other signs of sizelling of the cyto-
Oh
plasm are noticeable. TA) the contrary, at th2 temperatures 141441'
,.=e-lower than the limit of resistance, 4-4 swelling of the cyto-
plasm ?ccurs freque_nt13). These facts indicate that the caus4'?of
cell death from heat in different\zones temperaturf:;
coatd be quite different. In the zone of higher temperatures
04?10-,t
the leading cause remains the heat denaturitlg of the protoplasmic
proteins. During the continuous moderate heating, the death of the
cell occurs, presumably, as a result of a disruption of metabolism
)t
which leads to4mmds4 accumulation of toxic products (letergot
1936, 1937, 1960) and to the depletion of materials essential to
life (Lundeggr4 1930).
CONFIRM
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Similar disturbances of metabolism can lead tovazds the
Atinn
denaturims of the cell proteins (Nasonov and Alexandrov, 1940),
but this:secondary reaction is rather chemical in nature and /E
4-awisa not reperit heat denaturing.
7. The factors arlih_-.c:?.?,rrine heat resistance of cells
toury=caus,7_L-by
The cytoph
iological analysis of the reaction of the cells
fo-ttye. ki.1!5111,E:1!y heatishows that the result of this action
by which we judge the heat resistance of cells is determined by
-(;k6
several factors: (1) by resistance of protoplasmic proteins towax4IU
denaturing
resistpnce
(2) 11,- the
only after
during the
action of heat'2 ori',atlower temperatures, br the
of metabbIi6:' procesaes tat4z-zs a shift in temperature;
reparatory ability of the cell wit=ich dycur not
cessa..nox alto
the saeeetzion- of heating, butat lower temperatures
heating; (3) Wthe capacity of cells to increase heat
resistance/4n response to the injurious action of heattbry'Oeat
'hardenin .1 The participation of single factors in establishing
the heat resigtence depends-uponthe'teiprature,Ith dura'tioftof
hgating,' and the time paeeeid after the action.
Further investigations should be directed towards: (a) the
study of the physico-chemical and the biochemical mechanisms i51-4;01.11.
basis -those.
-rt
ave at the gWandat1o.a-of &iagle factors which determine the heat
resistance of the cells, and (b) the study of the ecological signifi-
cance of these factors; i. e., to determine the role which they play in
adaptation of plant organism to the surrounding temperature.
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In rega70 to the first point, our information is very limited.
We have some Idea about the character of ttv, injurious action of
heating, but we hardly know anything about the processes which are
connected with the reparatory action of cells and with the increlOse
vrt
ef their resistance. The task of our laboratory work is directed
at present to study the nature of these cell reactions. As far,
as the second point is concerned, the cytoecological phase of these
investigations is the subject of the following chapters.
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III. CYTOECOLOGICAL STUDIES OF HEAT RESISTANCE OF PLANT CELLS.
? 1. Problems of Cytoecology.
A continuous and manilold adaptation of organisms to the
surrounding medium takes place during the process of evolution.
The adaptive response towards the action of the medium occurs at
different levels of(the organization of the living matter: molecular,
cellular, organiic and superorgani=ie?X coenotic. For example,
the plant may secure the protection against the high temperature
of the medium by ??'Te selection of a proper coenosis, (such as
growth under protection of trees), by the seasonal periods of develop-
ment, by ,the increased transpiration, by the reactive increase cif
heat resistance of the cells during overheating, by tEZ. increased
stability of the cell proteins, etc.
The purpose of cytoecology consists in the study of molecular
and cellular adaptations properly; 1. e.
of the peculiarities of the
S,Ive
molecules and the cells which secure the
appearance of the adaptive
cuk,aetz-rs?
? results already at these lower levels of
the organization of nt livins
matter. -Without a profound investigation in the field of cytoecology
adaptation (4 organisms towards
wako Vt- it- ?
gimilartympossible the
a complete solution of the problem.of
the surrounding medium is impossible.
solution of practical/ important problems of acclimatization and
A
resistance against frost, heat, drought and salinity of both plans
and animals. The cytoecological investigations are directly connectma
A
with the cytophysiological studies of the effects of the surrounding
medium upon the cells.
One of the main forms of cellular adaptation (not the only
ct,
one) is the establishment of4relatiOnship between the level of cell
resistance to a certain factor and the degree:4-Df intensity of this
factor in the medium. ?
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warns
-31-
2. Heat resistance of cells and the temperature conditions
at which tile animals exist.
As has been pointed out above, one of the postulates of the
anon asso,sit,01:
denaturihg theory of injury and excitation is the admittance that
the native state of the protoplasmic proteins is unstable at the
temperatures compatible with the active life of the organism and is
maintained in a dynamic state by the energy of cell metabolism. This
postulate suggested, the existence of a relationship between the
Ike
surrounding temperature of an organism andAheat resistance of its
our-
proteins. Preliminary investigations carried by me with the cells
of the ciliated epithelium of a number of animals living at different
temperatures, (Alexandrov, 1952 b), and also by some other investi-
gators, (Battle, 1926; RunnstrOm, 1927, 1930, 19363 Patzl, 1933;
Adensamer, 1934), showed the existence of a relationship between
the heat resistance of the cells and the temperature conditions
under which the animals live. Later this postulate was confirmed
tvia t VAN," of
by Ushakov and his collaborators on a*vaat;zoolPgical material,
(Ushakov, 1955, 1956 a,1956 b, 1960 a, 1960 b; Svinkin, 1959;
Dzhamusova, 1960 a, 1960 b; Zhirmunskii, 1960; Zhirmunskii and Pisareva,
1960; Zhirmunskii and Tsu Li-Tsun, 1960, and others). (*)
wt refC
(*) These cytoecological investigations on animals are carried Am-
in the laboratory of evolutionary cytology (Docent B. P. Ushakov
in charge), and in the laboratory of cytology of protista (Prof.
Yu. I. Polyanskii in charge), both-am:at the Institute of Cytology,
Akademii Nauk, in Leningrad. '
CONFIDEEK
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? The above-mentioned regularity appears clearly when the
heat resistance of similar cells is compared in closely related
species which are living at different temperatures. For example,
in the frog Rana Ridibunda which lives in the south the cells
tLa-v,
of different tissues are more heat resistant as?eompere&mith
wt-re
those from Rana temporaria (the cells compared are those from
? cilated epithelium, spinal cord ganglia, epithelium of cornea,
A
, cartilage, muscle fibers, and spermatozoids). Special studies
6 cL i
d' Chad shown that at the foundation of the difference in heat resis-
ance of cells observed was the difference in stability of the
protoplasmic proteins towards denaturing action of heat.S.
(Alexandrov and Arronet, 1956; Panteleeva and Ushakov, 1956;
Brown, Nesvetaeva and Fizhenkol 1959; Kusakina, 1961).
Another important fact was established for a large number
of species of animals (sea urchins, certain worms, mollusks, fishes,
amphibia and reptiles), namely, that the heat resistance of the
cells of the specimen of the same species is very constant and-t.0
.:trst 4
el-oe-i-qper]--uptrthern or southern areas-of-origin
(Alexandrov, 1952; Alexandrov, Ouchakov et Poljansky, 1961;
Ushakov, 1955, 1956, 1958, 1960a; Ushakov and Zander, 1961;
Zhirmunskii and Pisareva, 1960'; Dzhamusova, 1960 c; Kusakina, 1960).
? V,
There is also no difference in heat resistance of cells in poikillothentio
animals collected both during the summer and the winter. This
1,441ca.tes
,s4644-fies also the stability of this characteristic within the
limit of the species (Alexandrov, 1952; Arronet, 1959; Shlyakhter,
T. A., 1959). (*)
(*) Contrary to these data N. A. Shlyakhter (1961) reports? the
?
heat resistance of muscles of,froLVRann temooraria)is somewhat
higher in tIas. summer ae,,-comparedwith77the winter. cki
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Cr. -17
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A remarkable stability of temperature limits during the
early stages of development of a series of marine invertebrates
was observed by Runnstr;m (1929, 1930, 1936). He considersAthe
limits ftemperature development as a constant physiological
characteristic of the species. This concerns also the heat
resistance of cell proteins (Ushakov, 1959 a, b; Ushakov and Kusakina, 1
1960), which apparently determines the range of temperatures
necessary for the normal embryogenesis.
10
In thm=spe-e4man of a cert=in spec1es,th3?sadaptation towsedia
? _
4Rteltemperaturel is achieved, as a rule, not through Zbe changes
in the heat resistance of protoplasmic proteins /but through some
mechanisms connected with the higher levels of organization,
mainly through behavior of the animal during seasonal and life
cycles. In the phylogenesis, however, the adaptation towards the
changed temperature conditions is achieved by establishment of
physiological races, or new species, with diferent resistance
of proteins towards the temperature factor. Therefore, the cell'S
-fay.
heat resistance, and the limits 6E'temperature $ ef development,
can serve the purposes of taxonomy, phylogenesis, and the genesis
10
of the whole fauna (Runnstrom 1930, 1936; Ushakov, 1959 a, b;
Zhirmunskii, 1960; Dzhamusova and Shapiro, 1960).
?resultsit? Ii
The materIalon?the animal cells shall be referred to
further when a comparison with similar data on the plant cells
will be made. This will give an opportunity to establish both
the general biological regularities:7114=WS-14- the differences
between the plant and animal cells.
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3. Heat resistance of cells and the temperature conditions
of the life of plants.
,he ecologies _.agnificance of the amount of heat resistance
A.
trtAt plant be cort71Ta7;44petween more or less closely-
related plants,/1 ist which live in entirely different temperature-
ck;
condition 4 ,end.7.1L7m..*T4 the same plants during different seasons
of the year. airing similar investigations two questions arise.
nrs,.
immediately: (1) 141-v.141(.1- resistance remaill0 the same in
different tissues and organs:7 (2) wL't?her it change/ with the
development oE the plant and with the growth and differentiation
of the tissuesE
C
a) The ?ieat resistance of cells from differentitissues.
The following two examples een illustrate the first
ct..;,-(A?troyt
yCTh
question: tho..-seeession of protoplasmic mot-ion after 5 min. of
heating occurs at the temperature of about 46? in the cells
from the leaf epidermis of cotton (var. 108f) -TBeress in the
$ V
f
Acells of thei ctE capsulesr/oat *50,0k (Alexandrov, 1956).
This difference apparently has some ecological significance.
AisensAtadt (1952) showed that during summer daysTlundethe
sunlight the temperature of the leaf tissues of cotton plants
(usually 1-5? lower than that of the air. This cooling is
achieved by high transpiration. At the same timelin the tissues
af-
of the capsule the temperature is 5-7 degrees higher thanli-dfr---
on the lighted side, and only 2-3 degrees on the shady side.
The second example is taken from four varieties of
Cess4t-:01,,
barley. 4D* secession of protoplasmic motion after 5 min.
_e4? Gic,.k. 4AI
heating took place at 45.5? in the.cells of epide: from the
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e.c sheath leaves whereas in the cells of the Spinules from the
A
awns, it occurred at 42.5 (Alexandrov, 1956). It is quite
possible that the different temperature regimof the tissues
a-re
is responsible for the difference. Burgerstein (1920) had de-
scribed in the awns of barley highly developed stomata and a
high rate of transpiration.
Th data presented show that the cells from different
tissues of the same plant may differ considerably in their heat
Because theanlmalrcells from different tissues
ct$ /71
show alsoidifferenpA heat resistance (Ushakov, 1960 a;
resistance.
a.
Rumyantsev, 1960), it is concluded that tile-comparison of4 heat
?
resistance in different organisms must be carried or in'quite
similar cells.
b) A connection between the heat resistance of Dlant cells
and the growth.
Special studies on the relationship between the heat
Aev.o u
resistance of cells and growth had been carried ,an in our laboratory
,q43
by Gorbani (1961, 1963). In the epidermis from Zebrine pendula
A
Schnizl.: Tradescantia fluminensis, Echeveria secunda Booth, and
a
Dactylis glomerata she had founddigreater sensitivity tozrida heatItt
/t
in the cells of the young growing leaves aA-rompaped,fwith the cells
of the leaves which had ceased to grow.
in sensitivity of the adult leaves taken
the same plant. For example, in Zebrine
There was no difference
1e_v
from differentlayer4s- of
pendula the curve,
heat resistance of the cells from the tip of the leaf of the 1st
internode (counting from the
loweitemperatures almost by
14
top) are-moved--iir-the--d-irection-of
two degrees, as?compared-with the curve
cc' similar cells taken from the leaves of the 3d and 4th inter-
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a
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-36-
nodes
(fig. 17). The cells of the leaf from the 3d internode
which had ceased to grow did not differ in their resistance from
the cells of the leaves taken from the lower internodes, such as
C.4.SSCA:tiol.1
the sixth. During the winter, after general seeeaslon of growth,
the difference in heat resistance of the cells taken from the
4
ct-r
tips of the leaves Erom-different internodes disappears. The
-eke
rise ob-the heat resistance of the cells from the leaves ofAupper
internodes accounts for this. In the growing leaves during the
summer months, the cells at the base of the leaf are less heat .
th ,t a-C
resistant as=compared=with those Lx*m the tip. This difference
fagained be explained by the higher rate of growth of the cells
Lo:
at the base as-empared-with that at LlIc tip of the leaf. In
the leaves of the lower internodes which had ceased growth, there
is no difference in heat resistance between the cells of the tip
and those from the base. In tile winter, after growth ceases in
the upper leaves, the heat resistance of the cells at the base
increases but it does not reach the level of that of the cells
,em the tip.
The relationship betean heat resistance and the
oxt growth?'hre still more clearly illustrated in the experiments
shown in Fig. 18 where there is a difference in heat resistancebeZwecx
Lseyr?
in the,Icei s Erom-epidemi of leaves of K9slanchoe blossfeldiana
taken from different internodes. Thus, the growing cells showed
a greater sensitivity toad t heating. The plant was treated with
hyd-k-Aavele
a:514,-reiit41=f maleic The growth had ceased and, at the same
time, the difference in heat resistance of the cells from the
upper and the lower leaves had disappeared. The leveling off
CONFIDENTIAL
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vi.et At,e. to a h "t40. Oc ?
of resistance took-pl&ee-by.,the increase t?Iresistance in the
(v_vo,i/ it cet(4tyt c,t)
upper leaves r6a-ohiti&fEelevel of the lower leaves.
A
Therefore, the growth cell cell elongation is
.a.5setc,,x-s":eti Ceet
a4eq.s.r) aannecterd with the inereoed sensitivity of the cells towsTrft
heat" In L plants taken at different developmental stages
there is no difference in resistance of the cells which had ceased
to grow (see for example fig. 19).
The facts discussed above show that for the comparison of
htz-twe0 `v% C ZIIS 0 ;?
,11.resistance /hea' it different plant., it is necessary to
take similar tissues which Liau ceaseu to grow. The dif-Zerence
in the developmental phases of the plant is less important.
c) The-neat resistance of nrotopla,-mic proteins and the
temperature conditions of life of plants.
The temperature conditions necessary for the existence of
the plants are dete=ined by the E-:*2:1C-!?,0f,e c*:: habitat, by
4
the selected microclimatic ov-lin,, and by the seasonal periods
of growth. However, the temperature of the surrounding medium
oc
does not always correspond to the temperature ab,which the plant
tissues Biebl (1950) is not quite correct** in stating that
the plants are helpless in regard to the temperature factor.
Thus, Lange (1959) has shown that in the desert of Njritania certain
plants (e. g.; Citrullus colocynthis) are capable of maintaining
the leaf temperature almost 129 lower than that of the air because
A
of increased transpiration. Similar data of Aisengiftadt (1952)
have been given above.
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tke
The problem oftirelationship between the resistance of
tf4e
plants totta=d.s 4action of high temperature and the temperature of
the habitat has not been extensively investigated (we do not consider
here the work on . thermophilic microorganisms). Sapper (1935)
h154 studied the heat resistance of entire plants belonging to
different families and had cmc to the conclusion that in spite
of certain exceptions there is a distinct connection between the
I