Plant Physiol.

(1967) 42, 76-88

Photosynthesis, Transpiration, Leaf Temperature, and Stomatal

Activity of Cotton Plants under Varying Water Potentials
J. E. Pallas, Jr., B. E. Michel, and D. G. Harris
Southern Piedmont Conservation Research Center, USDA, ARS, SWC, Watkinsville, Georgia
and
Department of Botany, University of Georgia, Athens, Georgia
Received April 22, 1966.

Summary. Cotton plants, Gossypiunt hirsutum L. were grown in a growth room

under incident radiation levels of 65, 35, and 17 Langleys per hour to determine the
effects of vapor pressure deficits (VPD's) of 2, 9, and 17 mm Hg at high soil water
potential, and the effects of decreasing soil water potential and reirrigation on
transpiration, leaf temperatture, stomatal activity, photosynthesis, and respiration at
a VPD of 9 mm Hg.
Transpiration was positively correlated with radiation level, air VPD and soil
water potential. Reirrigation following stress led to slow recovery, which may be
related to root damage occurring during stress. Leaf water potential decreased with,
but not as fast as, soil water potential.
Leaf temperature was usually positively correlated with light intensity and nega-
tively correlated with transpiration, air VPD, and soil water. At high soil water,
leaf temperatures ranged from a fraction of 1 to a few degrees above ambient,
except at medium and low light and a VPD of 19 mm Hg when they were slightly
below ambient, probably because of increased transpirational cooling. During low
soil water leaf temperatures as high as 3.4° above ambient were recorded. Reirriga-
tion reduced leaf temperature before appreciably increasing transpiration. The upper
leaf surface tended to be warmer than the lower at the beginning of the day and
when soil water was adequate; otherwise there was little difference or the lower
surface was warmer. This pattern seemed to reflect transpiration cooling and leaf
position effects.
Although stomata were more numerous in the lower than the upper epidermis,
most of the time a greater percentage of the upper were open. With sufficient soil
water present, stomata opened with light and closed with darkness. Fewer stomata
opened under low than high light intensity and under even moderate, as compared
with high soil water. It required several days following reirrigation for stomata to
regain original activity levels.
Apparent photosynthesis of cotton leaves occasionally oscillated with variable
amplittude and freqtlency. When soil water was adequate, photosynthesis was nearly
proportional to light intensity, with some indication of higher rates at higher VPD's.
As soil water decreased, photosynthesis first increased and then markedly decreased.
Following reirrigation, photosynthesis rapidly recovered.
Respiration was slowed moderately by decreasing soil water but increased before
watering. Respiration slowed with increasing leaf age only oIn leaves that were
previously tinder high light intensity.

Mlost micrometeorological studies presently deal lems involved has been presented by Collis-George
xvith changes in the whole or segments of pastures, (11). With the foregoing in mind, these stuidies
rangelands, forests, or ctultivated crops. In any of patterned after earlier stuidies on corn (29), were
these cases, it is the contribution of each plant directed toward understanding how young cotton
that eventually integrates into the response obtained plants react to environmental changes in htumidity
for the biota or commuinity. How some plants and soil water stress at several radiant energy
behave in relation to broad changes in the environ- Transpiration, photosynthesis, stomatal ac-
ment may be presently known in a general way, but
otur knowledge is incomplete, especially under con-
levels. leaftperature, anthesiratomatal-
tivity, leaf temperature, and respiration were eval-
ditioiis of environmental stress. A suirvey of prob- tuated.
76
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 PALLAS ET AL.-WATER POTENTIAL EFFECTS ON COTTON
Materials and Methods
Genterail. The test planit was Gossypiumn liirsut-
turnz L., Empire, adapted to the Southeastern United
States.
Procedures of handling the soil and plants prior
to experimentation were standardized as follows:
after Cecil sandy clay loam was passed through a
10-mesh screen and air-dried, 2.86 g of Krilitim
(Monsanto Chemical Company) (15) and 1 g of
dolomitic limestone per kg of soil were mixed with
the dry soil. The latter established a pH of 6.5.
-
WXater was added to the plastic limit and again the
soil was air-dried. jlust prior to uise, 2 g of 6-12-12
commercial fertilizer per kg of soil wac added.
WN hen the plants were 16 days old 0.1 g of NH4NOQ
per kg of soil was applied.
Pennington green-coated, acid-delinted seeds
were germinated in vermicuilite at 30°, selected and
transplanted 24 hours after sowing into 3.6 kg of
soil in asphalted 92-ouncc- cans. Sufficient water
was added to each can to approximate the -0.05
bar soil water matric potential determined from
(lesorption curves developed using pressulre mem-
brane (34) and tension table (22) apparatus.
Because these methods do not measure the other
major component of soil water potential, osmotic
potential, the latter is not incltuded in any of the
estimates of soil water potential presented in this
paper.
Forty-eight houirs after sowing, the plants were
transferred to the growth room and held at 250,
60 % relative htumidity, and 65 Langleys/hotur (1
Langley = 1 cal/cm2; all radiant energy values
reported were measured with a Beckman-Whitley
Total Radiometer, Model H 188-01) 1 for 14-hour
photoperiods; and at 20° and 90 % relative humidity
for 10-houir nyctoperiods. The cycle was repeated
uintil the plants were 30 days old. \Vhen weights
at the end of the day indicated average water
potential had dropped to or below -0.3 bar, suffi-
cient water was added to restore -0.05 bar. Aver-
age wind speed near the plants was 1800 mi/hour.
The lights were gradually raised to their re-
quiired intensity from 0600 to 0800 by increasing
voltage, and from 1800 to 2000 the intensity was
gradually reduced to off. What we have termed
flull, one-half, and one-foturth suinlight equiivalents
were obtained by uising 96 300w, 48 300w, and 48
150w incandescent lamps. To help cotunteract light
qulality effects on growth (14) becauise of an in-
creasing proportion of far-red as the growth roon
lights dimmed, the last 15. minuites of the photo-
- Trade names anid company names are included for
tlhe benefit of the reader and do not infer any endorse-
ment or preferential treatment of the product listed by the
Uniited States Departmcint of Agri:ultture.
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77
period was supplemented by tuirning oIn slim-line
cool-white flulorescents to suipply 6 Langleys/houir
of radiant energy.
Estimates by means of an Epply thermopile
with Schott filters indicated that of radiation in-
cident on tuirgid cotton leaves, 8 % was reflected,
80 % was absorbed, and 12 % was transmitted. Of
this, only 7 % of transmitted buit 55 % of reflected
was long-wave radiation (0.97 to 4 ,u). Abouit
70 % of the incoming long-wave radiation (6 %
of total radiation) was absorbed by leaves. All
valuies are near reported valLues (6, 32, 33).
The stiudy vwas divided into 2 parts. From the
31st throuigh the 36th day of age, the effects of
hulmidity at 3 radiation levels were stuidied. From
the 37th day of age to termination of the experi-
ment, the effects of soil water stress were stuidied
by means of an irrigation-drying-reirrigation cycle.
Julst before experimental day 1 (31st day of
age), 69 plants were selected for uiniformity, and
randomly divided into 3 groulps (A, B, and C).
In each grouip 10 plants were uised for measuirement
of transpiration, 2 for leaf temperatutre, 1 for
photosynthesis, 6 for resp.ration, and( 4 were des-
ignated for sacrifice or replacement of plants
damaged duiring handling. To provide equlal total
light exposuires prior to the start of part 2, the
grouips were shifted from bay to bay as in(dicated
in table I.
To minimize water stress effects while studxying
vapor pressulre deficit (V-PD) effects of the air on
transpiratioin and photosynthesis, all plants wvere
watered to approximately -0.05 bar at 2000 the
evening preceding days 1, 3, 5, aind 7. 'Matric
potential of -0.0n5 bar was selectedl in lieui of field
capacity because the latter cann't readily be es-
tablishedl in small containers. Aeratioln problems
were not anticipated at -0.0.5 bar.
After day 7 plants were not rewatered until
estimates indicated soil water matric potential hadl
reachedl -15; bars on an average in all contailners
Table I. Location of Plant Group (A-C) Durinq
Experiinicntation
All bays bad full sunlight equivalenit (65 Langleys/
hr) on days 2, 4, and 6. All other days the bays had
sunlight equivalent as follows: bay 1. full; bay 2. one-
half (35 Langleys/hr); and bay 3, one-fourth (17 Lang-
leys/hr). The ambient temperature w-as programmed
for 250 (measured in the outgoing air duct) and usually
maintained -v 10 throughout. The relative humidity was
programmed to be 60 %, except for day 1 (90 %) and
day) 5 (30 %) and seldom deviated as much as 3 %
Average air velocity was approximately 1800 m/hr
throughout the entire experiment.
Days Bay 1 Bay 2 Bay 3
1 and 2 A B C
3 and 4 C A B
5 and 6 B C
Ther-eafter A B C
78 PLANT PHYSIOLOGY

DAY 7 DAY 8
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See Legend Page 81
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 PALLAS ET AL.-WVATER POTENTIAL EFFECTS ON COTTON 79
DAY DAY 2

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See Legend Page 81

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 80 PLANT PHYSIOLOGY

DAY 13 DAY 14
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Copyright © 1967 American Society of Plant Biologists. All rights reserved.
 PALLAS ET AL. -\N'ATER POTENTIAL EFFECTS ON
in any 1 bay. Several days after rewatering, the
plants were harvested for fresh and dry weights
and undisturbed soil cores were taken for second
desorption curve determinations.
Transpiration. Beginning on day 1 and each
day thereafter, starting at 0600 and ending at 2000,
hourly weighings of each of 10 plants from each
bay were made. Daily leaf areas of all trans-
pirational plants were computed (width X length
X 0.77) (2) using measurements made from 1800
to 1900 hours. Transpiration is expressed as
g/dm2 of single surface leaf area.
Leaf Temperatutre. Temperature differences be-
tween both the upper and lower leaf surfaces and
the ambient air were measutred for 2 plants in
each bay. Each leaf thermocouple junction con-
tacted the leaf surface between major veins.
Earlier studies indicated insertion into leaves did
not increase sensitivity. The reference (ambient)
junctions were suspended in air about 15 cm from
the plant at the mean height of the plant.
.Stomatal Activity. Stomatal opening was de-
termined visually using a stomatal viewer (30) and
expressed as the percentage of stomata observed
to be open. Stomata on both the upper and lower
leaf surface of 2 of the most mature, unshaded
leaves were observed on each of 2 plants in each
bay, each hour. To minimize handling of individual
plants, 2 pairs of transpiration plants in each bay
were observed alternately.
Photosynthesis. Measurements of apparent pho-
tosynthesis were made on the most mature leaves
(exclusive of cotyledons). Because of equipment
limitations, only 1 cotton plant in each bay was
monitored. A 2-minute clock-activated stepping
switch with appropriate air pumps, solenoid valves,
filters, and flow meters permitted continuous pas-
sage of growth room air through leaf chambers
(3 liters/min, 76 m/hr) and automatic intermittent
sampling for analysis of growth room air, standard
gases, and leaf chamber air. The leaf chambers
were fashioned after Brun's model (10). Since
they dissipated 25 % of the incoming radiation, all
valtues listed in table I should be so reduced for
applicability to the photosynthetic stuldies. The
rates of net photosynthesis were obtained from
CO, differences between incoming and outgoing
chamber air. The detector-amplifier was a Model
15A Liston-Becker Infrared Analyzer. Another
L/B analyzer on a 2-mintute sampling sequence
meastured intermittently growth room air and ouit-
side air 8 feet above grotund level.
COTTON8 81
Soil Water Potential. Percent water by weight
(Ow) was computed by:
total wt - (tare wt of container +
Ow = 100 dry wt of soil + fresh wt of plant)
dry wt of soil
Estimates of the fresh weights of plants were
made by sacrificing 6 representative plants the night
before the first experimental day, 4 plants the
evening of day 6, and all plants immediately
upon termination of transpirational measuirements.
Growth during the experimental period was con-
sidered linear when adju1sting weights for calcu-
lation of soil water matric potentials. The latter
were derived from a water desorption curve from
an analysis of undisturbed soil samples obtained
by carefullIy cutting the metal containers away
from the soil of 4 transpiration plants from each
bay after the last transpiration weighing. For uin-
disturbed soil between -0.5 and -0.05 bar, the
Ow's averaged 2.5 lower than for distturbed soil.
Respiration. To obtain some estimate of the
effects of water stress on leaf tissue respiration,
a \Varburg constant voluime respirometer was em-
ployed. The water potentials of leaves were
measured using the Schardakow technique (35)).
At least 2 hours for equilibrating leaf discs with
sucrose solutions were determined to be necessary
for reliable measurements. Each flask's water po-
tential was reguilated to match that of the leaf
by adiusting KOH concentration (0.1 ml in cetiter
well, 0.5 ml in side arm) and adding 1.0 ml of the
appropriate concentration of sucrose solution to a
filter paper disc overlaid by a circle of plastic
screen upon which a freshly clut leaf disc was
placed. Flasks with side-attached center wells
were employed to permit use of intact leaf discs.
All operations and measurements were performed
uinder laboratory light (< 0.2 Langley/hr). Res-
piration was always measured at approximately the
same time of day (1400 to 1/700 hrs), thereby
minimizing possible diurnal variations (26).
Results
and Discussion
Tr-anspiration. The positive correlation between
transpiration and radiation (energy) level was
pronounced (fig 1, days 1, 3, and 5). At all 3
radiation levels transpiration rates at 30 % relative
humidity were nearly double those at 90 % (fig 1,
days 1 and 5). The evaporative demand is much

FIG 1 Measurements of percent open stomatz?, transpiration rates, and leaf-air temperature differences. For percent
open stomata, - - - - denotes upper epidermis of first pair of plants; - - - -, upper epidermis of second pair;
- - -, lower epidermis of first pair; and - . . - -, lower epidermis of second pair. For transpiration rates and
leaf-air temperature differences, . . .denotes one-fourth sunlight equivalent; - - -, one-half; and - , full, except
days 2, 4, and 6 when all plants were in full sunlight equivalent and curves refer to previous day's exposure.
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82
greater at the lower humidity. The ambient air
at 30 % relative humidity has a vapor pressure
(leficit (VPD) of approximately 17 mm Hg and a
water potential of approximately -1600 bars, as
conmpared with air at 90 % relative humidity, which
has corresponding values of 2 and -140.
.At all light levels anid all htumidities tinder high
soil water, cottonl trainspiration per UInit of leaf
surface was slightly greater than that which we
have reported for corn. Evaporation from a leaf
is regarded to be directly proportional to the v-apor
pressure difference between atmosphere and leaf,
andcl inversely proportional to resistance (9, 24).
Fewer stomata were open at low humidities than
at high humidities (fig 1, days 1 and 5).
As soil water potential decreased, transpiration
decreased, after first increasiing slightly at the
lower light intensities (fig 2a). The inability of
the plant to meet evaporative demand with de-
creasinlg soil water potential is demonstratedl by
the clecrease in transpiration at all radiation levels
at less than -1 bar matric potential. This is in-
terpreted to be due to inability of water transport
from soil to and through the plant to meet evap-
orative demand. Slatyer (38) has presented evi-
lence that the transpiration of Gossypiiumii barbai-
(lense L. decreases with decreases in soil water
potential.
The graphs (fig 2) reflect the increasing energy
requlire(d to move each increment, or specifically
each molecuile, of soil water into the atmosphere.
We have shown that a similar relationship exists
with the transpirationi of sorghLum and corn (29).
For corn the inflection point appeared to occur at
higher matric potentials with high evaporative de-
mclands: however, the smaller soil voltumes in those
stud(lies must be taken into consideration before
equatinig results. It has already been ahly demon-
strated that there is no magic figture of -15 bars
potential at which soil water is no longer available,
but rather there is a progressive decline in trans-
piration uintil the physical-chemical state of plant
Nx-ater makes it no longer capable of satisfying the
DAYS FROM INITIAL IRRIGATION
' 45 .
es 4 o
°O 3 5.
° 25 .
4 20 .
zs
z ID. I
C _
5
J2 os 3)1 0 0001 02 0304 060810
NEGATIVE SOIL W,A1FR POTENTIAL (BARS)
FIG. 2a-b. Average daily rates of transpliration at 3 light intenlsities as affected bx average daily ,soil water potenl-
tials during irrigation-drying-reirrigationi cycle. First day from initial irrigation. is experimental day 7. The 3 upper
horizonital scales shox. times corresixpnding to water potentials of abscissa. a) Fromii irrigationi tlhrough drying. b)
Follows-ing reirrigationl.
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PLANT PHYSIOLOGY
requirements for living cells.
After irrigation, transpirationl did not imme-
diately reattain the early high rates fouind at high
soil water (compare fig 2a with fig 2b). Al-
thouigh plants uinder high light did, those tinder
mediuim and low light did not, in the time allotted,
quiite reattain the transpiratioin rates of day 8
(day 2 from initial irrigatioin). The recoverv
rate was similar to recovery rates found for trans-
piration of pine btut not tomato (8). Correlated
with this is the observation (see fig 1, days 13
and on) that stomatal activity at any light intensity
on an average did not reattain the previotus highs
recorded on days 7, 8, or 9 (fig 1). Inability of
reattainiment may be relatecl to an impairment in the
absorptive capacity of the cotton roots which oc-
cuirrecl dutring the drouight. The fact that trans-
piration approached earlier rates in a couiple of
days leads us to stuspect that root damage was at
least partially of protoplasmic natuire.
.As soil water decreased, a slight decrease iin
transpiration, and at one-fouirth suinlight equivaleint
a rise in leaf temperatulre, were frequently notecl
in the late afternoon (see days 9 and 10). The
transpiration decline is consistent with, buit not as
apparent as, Skidmore aind Stone's observations oIn
nutrient-cultured cotton (37). At the same time,
stomatal observations in(dicated some closuire, possi-
bly explaining the redtuction. There is reported
an auLtoilomous rhythm associated with cotton
stomatal activity (13). \V:hether the stomata react
passively to water balance changes, and(l whether
such chainges are the resuilt of root impedance,
remains to be seen. It is well to recall that xvlem
exuidation also follows a flulctuiating diuirnal pattern
that seems to be impressed uipoIn it by the previous
history of the plants (21).
As the soil vater potential decreased, the water
potential of the cotton leaves decreased. A series
of representative valuies determined for vapor pres-
suire atljuistments in the \VNarburg flasks are liited
in table II. The matric water potential of the
soil exceeded the water potential of the leaves. In
DAYS FROM INITIAL IRRIGATION
E
Z
2
z
cr
20 30 4, 60180100150
01Di .. ..
02 03 04 .0 a601 02 0 304 06 08 10 20 3 40 80o30
.0
MEA4'IVF SOIL WATER POTENTIAL (BARS)
 PALL AS ET AL.-WN'ATER POTENTIAL EFFECTS ON COTTON
Table II. Soii Matric Potential (4j.) and Cotton Leaf
IIater Potentia-l (4ii) of Cotton Lcaz'cs fromii Plants
Under One-Fourth Sutnlight Equivalent Used in
Respiration Studs'
Day
qt*
9 - 0.05* -11-14*
13 --0.3 -14-17
16 - 8.0 -17-21
17 - 15.0 -22
17 -- 0.05 -11
* Valuies in bars.
the range of soil w-ater potential that this study
encompassed, leaf water potential of soil-grown
cottoin plants has been found to be several bars
less than the ater potential of the soil (38).
The final potential in the flow system is that of
the atmosphere, wvhich generally ranges from 100
to 3000 b)ars less than that of the plant. Present
techniiquies do not measture the true water potential
of stunlit leaves. Leaf water potential determine(d
Usinlg Schardakov or psychrometric techniquies is
biased strongly in favor of the water potential of
cells rather than that of substomatal cavities dutr-
ing transpiratioin. Dturiing the measuirement, trans-
pirationi is zero anid the gradienit between mesophyli
cell suirfaces and stomata disappears.
Leaf Terniperathire. Leaf temperatulre was highly
correlatel with tranispiration uinder most condlitions.
Huimiditx effects on cotton leaf temperatuire are
show-n in figuire 1, days 1, 3, and 5. 1iedian leaf
temperatuires w-ere 2.5, 1.5, and 0.50 higher than
ambient air temperatuire at high, meditum, and loxv
radiation levels, respectively, on day. 1 with high
huimidity. At high light intensity, days 2, 4, anfl
6, leaf temperatuires were mostly to 2° above
the ambient air temperatture in each bay. No leaf
temperatLures below ambient were recorded uintil
day 5 at low- relative humidity, (fig 1, day 5).
On this day, leaves at 1)oth meditum and low light
intensities dropped below the ambient temperatuires.
Leaves at high light tunder the higher VPD's (-
19 mg Hg) onl day 5 averaged 10 cooler than
(- 4 mm Hg) Ofl (lay 1. The higher transpiration,
and thuis increased latent heat exchange on day 5
compared with day 1, is shown in the markedly
lower l.af temperatture. In earlier stuidies with
corni, sorghuim, tomatoes, soybeans, and cottoin (28,
29) e consistently observed leaf temperatuires
above ambienit except with cotton plants. It ap-
pears, stibstantiatinig the wxork of Baker (3), that
there is a (lefinite tendenicy for illulminated cotton
leaves in air of high VPD to maintain a tempera-
ttre below% that of ambient air. The tendencv for
cotton leaves to be cooler tinder similar enviroIl-
mental condlitionis than the leaves of other crop
plants is not completelv explained by increases in
transpirationi (24). High advective heat exchange
may restult from openness of cottoni foliage. The
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83
latter might also affect radiationi absorption anid
reradiation balance. Ansari and Loomis (1) re-
ported that leaf temperatture of cotton dropped
below ambienit (3°) only utnder a very low artificial
light intensity (1 ft-c); whereas, Eaton and Beldlen
(16) have reported that leaf temperatulres on cotton
plants in the field tunder arid conditions frequiently
average(l as mulch as 3.5° below that of ambient.
Excellent positive correlations letween radiaint
energy aand leaf temperatture are recorded in the
literatture ( 1, 12, 20, 28, 30').
WVith decreases in soil water, transpirationi was
decreasing: meaniwhile, leaf temperattures rose. The
leaf temperatulre data oIn day 8, with high soil
water, indicate the averages of all sturfaces of all
leaves were 1.5, 0.5, and 0.10 above ambient at
high, oine-half, aind one-foturth suinlight eqtuivaleint,
respectively. Leaf temperattures of plants vn-!er
low soil water potential at the same radiant enirgy
levels dturing the 4 hoturs preceding irrigatioil were
3.4, 1.3, and 0.50 al)ove the ambient air temperatulre.
After irrigation, an almost immediate (1rop in leaf
temperatture w as noted at both high and(l m2alitim
light intensity, with returns to day 8 values several
days later. It is obvious from this anid other work
(12, 19, 41 ) that trainspiration certainlv cannot be
ignored in calcuilations of changes in leaf temper-
attire.
The decrease in leaf temperatuire after watering
cannot be explained by large increases in trans-
piration alone, as was the case with corn (29).
The positioin of the leaves appears to explaini the
record most adequiately. Cotton leaves failed to
show typical tuirgor-iindtuced movements as water
stress increased. In high light 3 (la s before irri-
gation, the lamiina failed to rise fii,1y duiring the
photoperiod. Couipled with th;s failuire was leaf
limpness. Both the lack of movemenit ain(d limpness
resulted in positioning of man) leaves to that of
minimum net capture of incoming radiation and
can be considered an adaptive mechanism similar
to leaf rolling in corn. Also on ('ay 12, iinder
high light, leaves failed to rise after irrigation;
therefore, very little radiation was directly im-
pinging on leaves. WVith less absorbed energy, the
leaf temperatuires woLuld be expected to be lower.
The same phenomenon was seen at medium light.
At low light levels, the phenomenon xvas observed
buit its magnittude was mulch less; not onlv was
the impinging radiation low, buit some changes in
leaf temperatuire may hav-e been below the sensi-
tivitv of the system.
Ouir earlier studies on corn indicated that a
1° leaf temperatulre difference existed between the
upper and lower epidermis. In these studies on
cotton more precise data were obtained and the
leaf temperatLure difference was fouind to vary
with radiant einergy and availability of soil water.
Under low light (fig 3) the temperatture gradient
was positive utntil the end of day 13, when it
became negative. After watering onl day 17, ani
84 PLANT PHYSIOLOGY

ences in the cuitictilar diffusion (28), the substoma-

0
40
Sunlight Equivlolnt
40 tal chamber configuration and stomatal iinumber, sinice
o- .0 . 130 there are less stomata on the tipper suirface of the
cotton leaf. Thuis, transpirational coolilng of the
2.0 .20 lower epidermis at this stage can be thouight of as
0
; ,0 exceeding that from the tipper suirface. \Vith an
inadequate suipply of soil water, a re(ldction in
10
.
0
transpiratioin tinder high light on (lay () (fig 1)
-OATERED

6 10 12 13 14 IS 16 IT is - -2 0 |SUNLIGHT EOJ1.t..Ek-
7 9 11
DAY
30
I.;

z -2.0
4.0 140
a0
1/2 S.Il2hght Equ.volnt

130 6
:E
CL
I(L
.0 220 ID

La d,
W 2
0
0n 0
lx
tS10 .-*,WTERED .
x
I.-
10 39
*0O 0
2:
0

-2.s _
7 9 10
.

11 12 13 14 5
_if16 1?
.' 0
-2
2
DAY HOURS

Zg 4.0 40
SUNLIGHT EOUIVALEN

OK
1/4 Sunlight Equivalent
.30 b 0--DAY FEFORE|
-- DAY OF tRRiGAT,ON
....- DAY AFTER
I IC a

20
el 0
1 25.0
01
aC
0l- 20.0
:i I
~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~
-2

8
r 10 11 12 13 14 15 16 17 is 2
05.II.O
io3d5 o.
___ aLEdE~~~~~~~~~~~~~~~~~~~"
DAY I
FIG. 3. Hourly averages of upper-lower leaf surface X S.
temperatures durinig irrigation-drying-reirrigation cycles.
: : : _
a 9 10 11 12 13 14 ID 16 is
upward trenid was seen. At one-half sunlight HOURS

eqtuivalent (fig 3) the lowest temperature differ-

ences between the 2 suirfaces were fouind( duiring
water suifficiency. A tendeincy existed for the uipper
suirface to be warmer early! in the day with the
lower stirface being warmer later in the day.
After watering, reattainiment of warmer tipper
a
stirfaces was eviidenced. At the beginning of the
day ulnder high radiant energy (fig 3), the tem-
perattire gradient was positive on days 7 and 8,
dropped on day 9, and remained negative uintil 0

rewatering on day 12, when again the tendency x

was to positive differentials.

We interpret the differences in leaf surface
temperatuires as follows: Under water stif ficiency
(leaf turgidl) the impinging radiation is screened
by the leaf and the lower epidermis receives the HOURS
least. Applicable in this stage of leaf water re- FIG. 4a. CO, concentration of the air and leaf photo-
lations are data of Van Bavel et al. (40) indicating synthesis under full sunlight equivalent, day 9; b-c)
that the lower epidermis, as compared with the Photosynthesis before and after irrigation. Sunlight equiv-
upper epidermis, has less resistance to water trans- alent irrigated on day 12; one-fourth stulight equivalent,
fer. The resistance is probably related to differ- clay 17.

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Copyright © 1967 American Society of Plant Biologists. All rights reserved.
 PALLAS ET AL.-WNATER POTENTIAL EFFECTS ON COTTON
occurred- concurrently, the lower leaf surface tem-
perattures began exceeding those of the tupper sur-
face, indicating increasing resistance to energy
exchange between that surface and its sturround-
ings. By dav 10 utnder the high radiation, the
water stress had catused nearly complete stomatal
closture, as well as near vertical positioning of
lamina (see above). The positioning favored an
increased entrapment of reradiated energy, the
stagnation of air within the foliage, and in some
instances permitted direct interception of radiation
from overhead by lower leaf sturfaces. All effec-
tively increased the lower surface temperatture.
After waterinig, a slow return to original conditions
was seen.
Stomtatal Activity. The effects of radiant
energ) and V-PD on stomatal activity during the
first 6 experimental days are summarized in figture
1. The percentages of stomata open at high radia-
tioni levels on davs 1, 3, and 5 were always sig-
nificantly greater than under low radiation, a
photoactive effect; in several instances the stomata
at the one-half light level behaved intermediately
but Inot predictably. Activity at low light levels
was significaintly lower than at higher light levels
on the suicceeding day. The indirect effect of
evaporative demand appears to be most noticeable
on day 5 for stomata under low light where the
ntumbers visibly open were fewer than at any time
dutring the hutmidity study but, in turn, rose to the
highest values observed on the following day at
full lig ht. Baker (4) surmised a stomatal effect
when at conistanit VIPD's transpiration increased
85
with increasing light intensity. The greater the
number of stomata open, the less is leaf resistance.
This partially explains any increases in transpiration
with increases in radiant energy.
In most instances throughout the studies a
greater percentage of stomata were open in the
upper epidermis than in the lower epidermis; this
is opposite to findings on corn in the growth room
(29), buit comparable to field studies on corn (28).
In the soil water potential study the greatest
percentages of open stomata were found during
water suifficiency, i.e., days 7 and 8. During this
period the effect of light intensity on the photo-
active operation was also noticeable; a significantly
greater number opened at high, fewer at medium,
and fewest at low light levels. No midday closure
was found; neither was there any indication that
stomata closed earlier on the upper surface than
those on the lower surface (36). As soil water
potential decreased, the number of stomata ob-
served to be open decreased at all light levels,
probably becatuse decreases in leaf water potential
eventually override the ability of guiard cells to
alter their turgidity. Considerable decreases in
number open occturred at high light when matric
potential was near -2 bars, at medium light near
-1 bar, and at low light near -0.1 bar. Boyer
has reported (7) that osmotic potential down to
-8.5 bars established with high NaCl concentra-
tions had no effect on stomatal resistance as meas-
tired with a porometer. Transpiration in his stuidy
was not reduced, probably reflecting the salt toler-
anit aspect of cotton water relations.

Table III. Average Leaf Apparent Photosynthetic Rates (P*) fromn 0800 to 1800 at Threc Light Intensities
Dar ** Light P* 4.in*** Light qjm*** Light P* 4Im***
1 12.3 I/2 10.5 7.9
2 . Full 14.8 Full 13.9
14.0 . .

. I /2 11.7 . . '/4 8.8 ..

4
1,,
16.0 Full 14.8 Full 12.6 .

5
P,,
16.3 ..
I/2 12.8 . . . '4 9.4
6 15.1 Fulll 14.4 Fulll 15.5
7 15.9 -0.03 /2 9.8 0.03 ~4 10.8 -
0.03
8 l,,
14.0 -0.1 9.5 - 0.04 l, 14.6 -
0.05
9 P,,
18.9 -0.23 15.2 - 0.05 14.8 -
0.08
10 t,, 14.6 -2.5 14.2 - 0.08 16.9 -
0.14
11 11.8 -4.0 15.1 - 0.13 P,, 16.6 -
0.26
12 11.1 - 6.0-0.03 14.2 - 0.3 1, 15.4 -
0.8
P,, 3.5
13 17.9 -0.05
01,
12.4 0.5 P,, 13.3 -

t,, 11.3 6.0

14 21.5 -0.11 11.9 -
1.9 1, -

15 ,,
9.2 -
3.3 7.6 -11.0
16 . PO, 6.7 - 55 P,, 5.1 -20.0
17t ..
10.8 80 P,, 8.9 -25.0-0.03
18 ,,P
.. P,, 10.4 -11.0 1, 10.8 -
0.9
19 "PI
.. ..
7.1 -15.0 11.0 -
3.3
20 ..
10.0 -20.0-0.05 ..

* Mg CO.,/dm2/hr.
** The relative humidity was programmed to be 60 %, except for day 1 (90 ,) and day 5 (30 %).
*** Best estimate of the soil matric potential (bars) at end of day and before irrigation. The first of 2 values is
that just before irrigation.
t The Ilant uinder one-fourth light was larger than the one under one-half light and required an earlier watering.
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Copyright © 1967 American Society of Plant Biologists. All rights reserved.
 PLANT PHYSIOLOGY

After irrigationi, stomatal activity resuimedl, al-
thotugh it took several days to reach original levels
of activity. Thuis, it was in(licated that the drouight
had not irreversibly affected the ability of the
gulard cells to operate. Decrease in plant water
potential in late afternoon is probably responsible
for the decreasedl number of stomata open dulring
this periodl. \Vhein other environmental factors
were not limitinig, a diuirlnal pattern of stomatal
activity was apparent, with photoactive openinig
occuirring shortly after lights came on and( closuire
after (lark.
Photosynthesis. On (laxvs 1, 3, and 5, radiant
energy can be seen in table III as a limiting factor
in photosynthesis. The relationship at any 1 hui-
midity was nearly linear between light intensity
and photosynthesis, which is consistenit with the
fiekl data of Baker (4).
A trend appears in table III on days 1, 3, and
5 at any light level, indlicating that the apparent
photosynthetic rate was related to the relative
htumidity of the air. The higher the V'PD of the
air, the greater the carbon fixation per uinit of
leaf area. This is not consistelnt w,% ith the resullts
of Bierhulizen and Slatyer ( 5), or Baker (4);
however, their studies were ruin at conisiderably
higher air temperatuires. Optimulm conditions for
photosynthesis of Deltapine cotton have been re-
ported to occulr aroin(l leaf temperatuires of 252
(26). The lowering of leaf temperature at the
higher VPD may thuis increase photosynithesis.
Throuighouit these stuidies the apparent photo-
synthetic rate oscillated (fig 4). Neither the
amplitude nor the frequency of oscillationi was
uniform. At least 2 reasons can be giv-en for the
oscillations: A) an effect on photosynthesis by
the concentration of CO, in the growth room air,
and( B) endogenous plant cycling phenomena.
Uinder high light intensity dluring days 2, 4,
an(l 6, all leaves of all grouips photosynthesized
at similar rates (table III), a fact that makes it
possible to place confideince not only in trefl(ls
appareint, buit also in suidden chanlges that occuLrred
(Iiring drawdown and followilng irrigation. The
effect of soil water stress oni photosynthesis is
'deilt in table IIT and figulre 4.
Dturing the drawdown perio(I a very initerestillg
tren(l occulrre(I at all light intenisities, a rise and then

1/2 SUNLIGHT EOUIVALENT

Loaf No Not loogcted 1-go'te
S a-A
6 _
7 -t U

t,
E
i7--

0N
a I,

b
, L
2 3 4 5 6 7 8 9 10
DAYS FROM INITIAL IRRIGATION DAYS FROM INITIAL IRRIGATION

U J VT T El' T
4 SUN9LlG,T FOUJIVALEN' Nof I" g5r*o: ,, gV64
Leaf Nto %o2 i-godte -.gated 3.0~ SUNLIGHT EQUIVALENT
4 1/2 SUNLIGHT EOUIVALENT ---
5 1/4 SUNLIGHT EQUIVALENT *-a

SC

6
102 C

z.-
0 -
r 2.0
O IC O OF0
C .zd
2 3 4 5 6 7 8 9 10 11 2 3 4 5 6 7 £ c 10 11
DAYS FROM INITIAL IRRIGATION DAYS FROM INITIAL IRRICATI-,/

Fic. 5a-c. Average respiration rates of individual leaves as measuire(l unider subdued light. Soil water potenitial
decreases from left to riglit (see fig 2 and tables II and III); d) averages of all leaves. First day froimi initial
irrigation is explerimeintal dav 7.
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Copyright © 1967 American Society of Plant Biologists. All rights reserved.
 PALLAS ET AL. -WATER POTENTIAL
a fall in photosynthesis. The high rates of photo-
synthesis occurred several days after the beginning
of the drawdown cycle. El-Sharkawy, et al. (17)
noted higher photosynthetic rates of cotton in dry
soil as compared to wet.
At the lowest soil water potential imposed,
photosynthesis rates decreased to 42 % under full,
66 % tinder one-half, and 52 % at one-foturth sun-
light equivalent of their previotus maximum rates.
This correlates well with the report (18) that
visibly wilted cotton leaves maintained relatively
high photosynthetic rates. Corn photosynthesis
dropped to the compensation point at similar low
water potentials (29). Low plant water potential
(-12 to -15 bars) induced by low soil water
has also been found to stop pine and( tomato
photosynthesis (8). Continuied cotton photosyn-
thesis under water stress does help explain the
findings of Phillips (31) that irrigation at the
25 % available soil water level was as effective
in increasing yields of cotton as irrigation at the
50 % available soil water level. The latter level
is most frequently used in field irrigation for sus-
taining yields. WVithin several houirs after rewater-
ing, an increase in photosynthesis was detected.
Under full stunlight equivalent, it eventually ex-
ceeded its initial high value. However, at one-
foturth light, the increase 2 days after atering
was only 65 % of initial photosynthesis.
An obviouls correlation existed at all light levels
between visible stomatal closuire and decreases in
photosynthesis (fig 1 and table III) suipporting
El-Sharkawy and Hesketh's findings (18); how-
ever, it is not likely that stomatal closuire was
completely responsible for decreases in photosyn-
thesis.
Respiration. Similar to reports on other species
(20, 39) as well as on cotton (25), older leaves
from plants under full sunlight equivalent had
lower respiration rates than younger leaves, al-
thouigh not unequiivocall) so (fig 5a). A similar
trend was not apparent with leaf samples from
one-half (fig 5b) and one-fouirth (fig 5c) light.
Effects of age were also reflected by plants irri-
gated to near field capacity throuighotut the sttudy
and measured on days 6, 9, and 11 from initial
irrigation (fig 5a-d). Some decrease occurred in
fulll sunlight eqtuivalent; less in one-half; and little
change in one-fourth.
At all light levels perceptible drops followed
by decided rises in respiration with decreases in
soil water potential were detected (fig .;a-d). Av-
erage decreases were 40, 33, and 20 % at fuill,
one-half, and one-fouirth sunlight equivalent, re-
spectively. Boyer's work (7) indicated a 25 %
drop in cotton respiration with lowering of osmotic
potential in cultu re solution to -8.5 bars. The
increased rates of respiration as soil water po-
tentials continued to decrease are not uinusual
(8,23).
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Copyright © 1967 American Society of Plant Biologists. All rights reserved.
EFFECTS ON COTTON /
Acknowledgments
Supported in part by the Meteorology Departmenit,
United States Army Electronics Research anid Develop-
ment Activity, Fort Huachuca, Arizona.
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