Published January, 1986
Leaf Photosynthesis and its Correlation with Leaf Area1
A. S. Bhagsari and R. H. Brown2
ABSTRACT
Comparisons among cultivars of leaf CO2 exchange rate (CER)
based on leaf area may be confounded by leaf characteristics
not usually accounted for in such comparisons. One such con-
founding characteristic appears to be leaf size. Correlation coef-
ficients between leaf CER and leaf area (LA) were determined
for genotypes of peanut (Arachis hypogaea L.), soybean [Glycine
max (L.) Merr.J, and sweet potato (Ipomoea batatas L.) and were
compared with correlations for other crops reported in the lit-
erature. There were negative correlations between CER and LA
in all crops examined. Significant negative correlations between
LA and dark respiration were observed for wheat (Triticum sp.),
alfalfa, (Medicago saliva L.), and peanut. Leaf area was nega-
tively correlated with transpiration per unit LA and with leaf
conductance for soybean. The relationship between leaf size and
specific leaf weight was inconsistent in most crops. Comparisons
of CER for genotypes having different leaf sizes may not rep-
resent the inherent differences in photosynthetic potential. Neg-
ative correlations between LA and CER may be one of the causes
for the absence of consistent relationships between CER and
yield.
Additional index words: CO, exchange rate, Dark respiration,
Specific leaf weight.
M ANY attempts have been made to relate CO2
exchange rate (CER) to yield of crop plants
and to select plants for high CER in order to improve
yield. In most comparisons of crop genotypes, the
relationship of leaf CER to yield was found to be poor
(3, 4, 11), probably because CER measurements in-
adequately represented the total seasonal photosyn-
thesis of the plant canopies (32). Differences in CER
among genotypes have been shown foe several crop
species and nave been compared to differences in
various leaf anatomical, physiological, and biochem-
ical characteristics. In some cases close correlations
have been observed and in others there were no cor-
relations. Specific leaf weight (SLW) is one leaf char-
acteristic most often correlated (positively) with CER
(10, 11, 12, 18), but in some studies the two were not
related (17, 29).
Leaf thickness, a major component of SLW, was
positively correlated with CER in soybean [Glycine
max (L.) Merr.] (12) and alfalfa (Medicago saliva L.)
(10), but in cotton (Gossypium sp.), leaf thickness was
negatively correlated with CER. This negative cor-
relation may have been caused by the diversity of
Gossypium species included in the experiment (15).
Several studies have indicated that leaf CER based
on area was inversely related to leaf size. Compari-
sons of diploid wheat (Triticum, Aegilops) species with
modern tetraploid and hexaploid cultivars show that
in evolution of wheat, the leaves have become larger,
but CER on a leaf area basis has decreased (16, 21).
Significant negative correlations have been observed
between CER and leaf area (LA) in genotype com-
parisons of alfalfa (10), soybean (18), rice (Oryza sativa
L.) (24), and perennial ryegrass (Lolium perenne L.)
(30).
On the basis of relationships reported in the lit-
erature between LA and leaf N content for soybean
(18) and wheat (Triticum aestivum L.) (21), Hesketh et
al. (18) postulated a "dilution" hypothesis implying
that the photosynthetic pigments or enzymes per unit
LA become fewer as the area per leaf increases. This
event would lead to lower CER for large leaves com-
pared with small leaves. However, in experiments with
soybean, Hesketh et al. (18) failed to confirm that the
negative relationship between CER and LA was due
to dilution of specific enzymes or total protein.
Other gas exchange traits such as transpiration,
dark respiration, and photorespiration measure-
ments may be related to LA. Few reports contain
information on these relationships even when the rel-
evant data were collected. The influence of LA on
1
Contribution of the Agric. Res. Stn., Fort Valley State College,
Fort Valley, GA., and Agronomy Dep., Univ. of Georgia, Athens,
GA. Received 11 Mar. T985.
2
Associate professor, Agric. Res. Stn., Fort Valley State College,
Fort Valley, GA. 31030, and professor, Agronomy Dep., Univ. of
Georgia, Athens, GA. 30602.
128 CROP SCIENCE, VOL. 26, JANUARY-FEBRUARY
1986

1979 and 1980. These genotypes showed a wide range in

bgas exchange traits and other leaf characteristics may
e important in assessing the relationship of CER to
yield.
The objectives of this report are to determine if
negative correlations between LA and photosynthesis
are common across plant species, and to examine
other characteristics which may account for such a
relationship.
MATERIALS AND METHODS
The correlations presented in this paper are derived from
both published and unpublished sources. Experiments over
the period from 1971 to 1981 in which the authors have
been involved are the source for some of the data for the
CERvs. LA correlations for peanut (6, 7) soybean (1,
and sweet potato [Ipomoea batata (L.)] (4, 8). These exper-
iments were conducted to study genotypic differences in
CERand other physiological traits. Although LA was mea-
sured in these experiments, its relationship with CERwas
not examined.
Methods for plant culture and physiological measure-
ments are detailed in the reports from which data were
taken. Only a brief description of the experimental material
and conditions is given below. Twenty four peanut cultivars
collected from 17 different major peanut growing countries
were used in the correlations (6). Six wild species of peanut,
which were included in reference 6, were not used in cal-
culating correlation coefficients. Soybean germplasm con-
sisted of 16 determinate genotypes, four each from ma-
turity groups V through VIII. Two soybean experiments
were conducted during the fall of 1974 under greenhouse
conditions (5). The same set of 16 genotypes was used
a 1975 field experiment to determine CER during vege-
tative, flowering, and at rapid pod-fill stage (1). Field plants
were grown under irrigation, following standard cultural
practices. Carbon exchange rates were determined by pass-
ing compressed air containing 330 #L L-~ COs over young,
fully expanded leaves from the top of the canopy in full
sunlight between 1100 and 1500 h (EST).
Forty sweet potato genotypes, consisting of important
edible-type cultivars, industrial types, and new experimen-
tal selections, were obtained from Research Stations at
Chase, LA., and Tuskegee Institute, AL., and the Coastal
Plain Experiment Station, Tifton, GA. Photosynthesis was
determined under greenhouse and field conditions during
LA and shape. Somegenotypes had nugget (okra-leaf shape)
type leaves. The youngest fully-expanded leaves near the
tip end of the branch, fully exposed to sunlight, were used
for measuring CER. Other details were previously de-
scribed (4, 8). Genotype means in each experiment were
used for regression analysis of CERon LA in the peanut,
soybean, and sweet potato studies described above.
The data for cassava (Manihot esculenta Crantz) presented
in Table 1 are from unpublished experiments conducted
during the summer of 1982 under field conditions at the
Agricultural Research Station, Fort Valley State College.
The seven cultivars used in this study were obtained from
the Homestead Experiment Station, a substation of the
Florida Agricultural Experiment Stations. The cuttings,
obtained in mid-February 1982, were planted in pots in a
greenhouse and were transplanted, 31 Mar. 1982, in the
field on ridges 1.2 m apart, with 1.2 mspace between plants.
Fully-expanded leaves, 15 to 20 days old, from the top of
the canopy were used to determine CERin full sunlight in
mid-September. Four measurements were made for each
cultivar. The method of CERdetermination was similar to
the one already described for sweet potato (4, 8). For cas-
sava, single CERdeterminations were used for regression
analysis of CERon LA.
In addition to the cultivar comparisons described above,
relationships between CER and LA within genotypes were
examined in one peanut and two sweet potato genotypes
(unpublished data). A peanut genotype from Tanzania, Af-
rica (USDAPI no. 149263), was grown in large pots in the
summer of 1971 as described for the genotype comparisons
of Bhagsari and Brown (6). Carbon dioxide exchange rate
was measured at about 1300 h (EST) on 19 days. Fully-
expanded leaves from the tops of the plants were selected
and measured. The original purpose of the measurements
was to monitor changes over time in CERof a single geno-
type during the time when 24 genotypes were being com-
pared. (6).
Correlations between CER and LA were determined for
two sweet potato advanced selections from the breeding
program at the Coastal Plain Experiment Station, Tifton,
GA(75-96-1 and 73-61-1). These plants had okra-type
leaves, the lobes of which were larger but similar in shape
to peanut leaflets. For each selection, 12 fully-expanded
leaves, 15 to 20 days old and differing in size, were selected
near the tips of the branches for CERmeasurements. The

Table1. Correlationcoefficients (r) for the relationships of leaf area (LA) with C02exchangerates andspecific leaf weight(SLW),
rangein leaf area for cultivar comparisons of peanut, soybean,sweetpotato, andcassava.
Exp. no. LAvs. CER n~ CERequation.~ LAvs. SLW
Ran~in LACrop, location, and year Reference
r a b r acm
1 -0.61"* 24 31.0 28.0 -0.002 19- 39 Peanut,greenhouse,1971 6
2 -0.79** 16 35.5 35.0 -0.81"* 33- 54 Soybean,greenhouse, 1974(Exp.1) 5
3 -0.94*** 16 32.5 20.0 +0.25 41- 77 Soybean,greenhouse, 1974(Exp.2) 5
4 - 0.58* 16 29.7 10.0 + 0.62* 54-177 Soybean
vegetative,field, 1975" 2
5 -0.60* 16 30.2 12.0 +0.35 60- 97 Soybean,pod-formation, field, 1975 2
6 - 0.60* 16 29.7 12.0 + 0.22 46-106 Soybean,
pod-fill, field, 1975 2
7 -0.22 17 15.1 2.5 +0.05 48- 95 Sweetpotato,LA,greenhouse, 1979 4
8 -0.51" 19 17.5 4.4 -0.17 50-143 Sweetpotato, Tifton, greenhouse, 1979 8
9 -0.29 20 18.7 2.5 -0.24 76-172 Sweetpotato,Tiftonfield, 1979 8
10 - 0.63** 16 25.4 3.8 + 0.03 69-188 Sweetpotato,Tifton,field, July1980 8
11 -0.63** 16 26.3 5.7 -0.26 89-215 Sweetpotato, Tifton, field, August 1980 8
12 -0.81"** 16 23.7 4.4 -0.12 100-208 Sweetpotato,Tifton, field, September 1980 8
13 -0.71"* 15 25.0 5.0 -0.09 71-181 Sweetpotato,LA,field, July1980 4
14 -0.75** 15 29.0 6.9 -0.20 94-192 Sweetpotato,LA,field, August 1980 4
15 -0.77** 15 25.0 5.0 -0.03 62-186 Sweetpotato,LA,field, September 1980 4
16 -0.39* 7 17.7 2.5 +0.28 108-298 Cassava,field, September 1982 unpublished
data
*,**,***Significantat 0.05,0.01, and0.001probabilitylevels, respectively.
J" n -- numberof cultivars compare&
:~ Regressioncoefficientsfor the linear regressionCER = a - b (LA),withCER in t~nolCO,m-s- andLAin mm.
 BHAGSARI& BROWN:CORRELATIONOF PHOTOSYNTHESISAND LEAF AREA 129

plants were grown in the field during the summer of 1979 to -0.81 (Table 1, Exp. 7 to 15). Coefficients were
and 1980, and CERwas measured as described earlier (4, significant in seven out of nine comparisons. Like
8). For regression analysis of CERon LAwithin the peanut soybeans, LA for sweet potato was larger for field-
and sweet potato genotypes individual leaf data were used. than for greenhouse-grown plants.
Remaining sources of data for correlation of CER with The r for the relationship between CER and LA
LA(Table 2) were published reports of others. Correlation for cassava (Table 1) was also negative (-0.39).
coefficients from Evans and Dunstone (16) are based on the range in leaf size for cassava was greater than for
data obtained from their Table 2. For Khan and Tsunoda’s
peanut, soybean, and sweet potato.
wheat data (22), LA measurements from their Table 2 for For comparisons of LA and CER within the peanut
December 1968 and January 1969 and the corresponding
highest estimated CERvalues for the respective genotypes genotype, PI no. 149268, LA varied from 29 to 48
cm~ and CER ranged from 16.0 to 26.9 #mol CO2
from their Fig. 1 and 2 were used to calculate correlation
coefficients. For barley (Hordeumvulgate L.), CERand flag m-~ s-~. The r for the linear regression of CER on
leaf areas of lines selected for small and large leaves were LA was --0.59 (P < 0.01) for the 19 CER deter-
taken from Table 1 of Reference 3 to calculate r. Calcu- minations. The ranges in LA and CER for the sweet
lations were made separately for the large and small leaf potato selection 75-96-1 were 74 to 126 cm~ and 18.0
lines. Correlation coefficients for CERand LAfor other to 33.2 #tool CO~. . m-~ s -L, respectivel, y and the r was
crops are presented as reported in literature. -0.43 (not s~gnlficant). For selection 73-61-1,
The gas exchange reported in this paper is for single varied from 56 to 136 cm~ and CER from 14.1 to
leaves and, in most cases, was measured in acrylic plastic 28.8 t~mol CO~ m-~ s -~ with an r of --0.69 (P
photosynthetic chambers. The dimensions of photosyn- 0.05). The negative correlation coefficients for pea-
thesis chambers varied according to leaf size and shape. nut and sweet potato, given above, show that larger
Leaf environmental conditions during CER measurements leaves are associated with lower CER even within
are described in the relevant literature cited. In most stud- genotypes.
ies represented by data presented in this paper, CO2con- No s~gnificant correlation was found for relation-
centrations entering the photosynthesis chamber varied be- ships between LA and SLW for peanut and sweet
tween 300 and 340 I~L L-~ and temperature varied from potato (Table 1). The range of r for the regression
30 to 35°C for warm season crops and 15 to 25°C for cool of SLW on LA was +0.62 to --0.81 for soybeans
season crops. Photosynthesis was generally measured under and only the two extreme values were significant.
saturating light conditions. Correlation coefficients for the CER vs. LA rela-
Correlations for relationships between LAand leaf traits
tionships reported or calculated from published data
other than CERare presented for some crops in Table 3.
Dark respiration rates and SLWwere estimated for wheat of others are presented in Table 2 for several crops.
from Fig. 10 and 12 of Reference 21, respectively, and were All r-values were negative and were significant except
correlated with leaf area values in Table 1 of Reference in one experiment each for barley (3) and wheat (22).
21. For soybean (5) and peanut (7), LA was correlated The range ofr was from -0.45 to -0.87. In barley,
transpiration and leaf conductance (boundary layer and sto- 10 lines were selected for large leaves and 10 for
mata) for soybeans and with dark respiration rates for pea- small leaves from two populations generated by hy-
nut. bridizing small and large leaf parents (3). Although
Along with linear regressions, logarithmic, exponential, the mean CER did not differ for the small and large
and power regressions were also performed. No other leaf lines, CER and LA were negatively correlated
regression analysis consistently resulted in higher correla- within the large and small groups (Table 2).
tion coefficient values than linear regression. For example, Correlation coefficients for relationships between
r by linear regression for vegetative stage of soybean (Table LA and other leaf traits from the published literature
1, Exp. 4) was --0.58. For the same data, r by logarithmic, are presented in Table 3. Leaf area showed signifi-
exponential, and power regression were -0.62, -0.56, cant negative correlations with dark respiration for
and -0.59, respectively. Thus, only linear regression anal- peanut (7), alfalfa (10), and wheat (22). Transpiration
ysis data are presented.
Table2. Correlationcoefficients It) for leaf area vs. photosyn-
thesis (CER}frompublishedreports.
RESULTS
Plant Reference r
Peanut CER showed a significant negative corre-
lation (r = -0.61) with LA, which varied from Wheat 16 -0.70*
Wheat(December 1968) 22 - 0.45 NS
to 39 cm2 (Table 1, Exp. 1). Correlation coefficients (January1969) - 0.66**
for the relationship between CER and LA for soy- WheatJ" 1 - 0.87***
bean experiments ranged from -0.58 to -0.94. Barley(Smalllcaves)~: 3 - 0.71*
(Largeleaves) - 0.62 NS
Correlation coefficients for field-grown soybeans at Rice~ 24 -0.87**
three stages of growth, vegetative, pod-formation, and Soybeanf 18 - 0.66**
pod-fill stages, were similar (r ~ -0.60) as were the Alfalfa~" 10 -0.50**
Nicotianasanderaehort. 25 -0.66**
equations describing the relationships (Table 1, Exp. Loliumperenne{Lowlightl§ 30 -0.54*
4 to 6). The range in LA for greenhouse-grown plants (Highlight) -0.80**
was lower than that of field-grown soybeans at all *,**Significantat 0.05and0.01probability levels, respectively;NS--not
stages of growth and regression slopes and r-values significant.
were more negative (Table 1, Exp. 2 and 3). ~ r-valuesfor thesecropsare as reported
in corresponding
references.
The correlations for the relationship between CER ~:Tenlines selectedfor smallleavesand10 lines selectedfor large
leavesfromtwopopulations.
and LA o£sweet potato genotypes in greenhouse and § Photosynthesis wasdetermined manometricallyat 15°Cat light in-
field experiments were negative; r ranged from --0.22 tensitiesof 400and3000ft-c.
130 CROP SCIENCE, VOL. 26,
Table 3. Correlations of leaf area with other leaf physiological
characteristics.
Correlated
Plant Reference characteristic
Rice~ 24 Soluble protein~
V(hcat 22 Darkrespiration:~
December1968
Dark respiration~
January 1969
TranspirationS/
January 1969
SLW,December 1968
SLW,January 1969
Alfalfa~f 10 Dark respiration~
Soybean 5 Transpiration~
Leaf conductance
Peanut Dark respiration~
7 -0.56*
*,** Significant at 0.05 and 0.01 probabilitylevels, respectively; NS--not
significant.
J"~valuesfor these crops are as reported in corresponding
references.
:~Expressed on a leaf area basis.
of soybean (5) was negatively correlated with LA
= --0.94) but no relationship was observed between
LA and transpiration for wheat (22). Leaf conduct-
ance was positively correlated with LA of soybean
(5). In wheat (22), SLWwas negatively correlated
with LA, with a significant r-value in 1 out of 2 yrs.
Correlation coefficients for the relationship be-
tween CO~in the photosynthesis chamber (average
of entering and exiting concentrations) and CERwere
determined for Exp. 1 to 16 in Table 1. The r-value
ranged from -0.50 to +0.28 and were not signifi-
cant except for cassava (r = --0.50, P < 0.05), in-
dicating that differential depletion of CO~due to leaf
size was not the cause of the negative relationship
observed between CER and LA.
DISCUSSION
Selection of genotypes with high CERis important
for improving crop yields, but screening for high CER
may be confounded by interactions with other leaf
traits and environmental factors. From a survey of
literature, Elmore (14) concluded that true genotypic
comparisons of CER were confounded as researchers
may be measuring the adaptive responses of the geno-
types to light, nutrient stress, temperature, or other
known and unknown environmental factors.
Leaf area may also be a trait which, if variable,
confounds the comparisons of CER among geno-
types. The consistency and stability of correlations
between CERand biological and economic yield may
be affected when genotypes under study have differ-
ent leaf sizes or when leaves of comparable area are
not selected for comparison.
From comparisons of CER among genotypes (Ta-
bles 1, 2), and comparisons within genotypes of pea-
nut and sweet potato, it is evident that LA and CER
are negatively correlated. The causes for the negative
correlation between leaf size and CERare not known
but the "dilution" hypothesis of Hesketh et al. (18)
does not appear to be involved, since no relationship
between LA and SLWwas found for alfalfa (27) nor
in most experiments included in this report (Tables
1, 3). In addition, Hesketh et al. (18) found weak
Correlations between LA and ribulose bisphosphate
carboxylase activity (r = --0.41), indicating not much
dilution of this enzymein large soybean leaves. Lack
JANUARY-FEBRUARY1986
of dilution is also indicated in the work with wheat
species (2), where a high correlation (r = -0.87)
found between LA and CER on a leaf area basis.
r WhenCERwas expressed on a chlorophyll basis, the
-0.90** correlation calculated from data in (2) was still neg-
- 0.66** ative (r = --0.58, P < 0.05). It can be calculated
from data of Jellings and Leech (19) that the chlo-
- 0.56"
roplast numbers (in mesophyll) per unit leaf width
- 0.05 NS increased rather than decreased with increased leaf
size in tetraploid and hexaploid wheats even though
-0.53*
-0.42 NS mesophyll cell numbers decreased.
- 0.46** Sometimes large leaves are thinner than smaller
- 0.94** leaves as indicated by the negative correlation (r
- 0.56*
-0.75). between thickness and area per leaflet for
alfalfa (10) and, in this case, there was a positive cor-
relation between leaf thickness and CER.It is possible
that negative correlations between LA and CER (Ta-
bles 1 and 2) resulted from small leaves being thicker
than large leaves. However, the lack of correlation
between LA and SLW(Table 1) minimizes this pos-
sibility. What may be involved is the tendency for
larger leaves to have larger mesophyll cells. In com-
parisons of alfalfa leaves of different sizes due to po-
sition, Turrell (28) found larger leaves to be thicker,
et they contained fewer mesophyll cells per unit area
than smaller leaves, thus, lnd~caung larger cells ~n
large leaves. Moderncultivars of tetraploid and hex-
aploid wheat have been shown to possess larger leaves
and larger mesophyll cells than ancient diploid spe-
cies (2, 13). If cultivars of the same species with large
leaves also have larger mesophyll cells, then CERmay
be reduced, since Wilson and Cooper (30), and Austin
et al. (2) found a strong negative correlation between
photosynthesis and mesophyll cell size. Wilson and
Cooper (30) attributed the benefits of smaller me-
sophyll cells on CERto an increase in exposed me-
sophyll surfaces for CO2uptake. In Turrell’s (28)
analysis of large and small alfalfa leaves, however, he
found the ratio of exposed internal cell surface to
leaf area to be positively rather than negatively re-
lated to leaf size, mainly because larger leaves had
thicker mesophyll. Thus, causes for effects of cell size
on photosynthesis are uncertain.
It is knownthat with increasing ploidy level, cell
size often increases making the leaves and other plant
parts larger. From a comparison of CER of diploid
and tetraploid wild wheat species and cultivated wheat
at three ploidy levels, it has been found that in the
process of evolution, the LA has increased with a
corresponding decrease in CERper unit leaf area (2,
16). Correlations between ploidy level and CER do
not always occur because wild tetraploids and hex-
aploids synthesized from wild species showed higher
CERthan cultivated diploid species (21, 22). In ad-
dition, increasing ploidy level in alfalfa did not de-
crease CER even though LA was increased (26).
One possibility for the inverse relationship be-
tween CER and LA could be lower CO~ concentra-
tion in the leaf chamber for large than small leaves
since large leaves would cause greater depletion of
CO2if constant flow rates were used. However, in
experiments using high flow rates to minimize CO2
depletion in the leaf chamber (18), or utilizing var-
iable flow rates to give maximumCERfor each geno-
type (22), LAstill showedan inverse relationship with
 BHAGSARI & BROWN: CORRELATION OF PHOTOSYNTHESIS AND LEAF AREA 131

CER (Table 2). In the experiments reported in Table

1, no relationship was found between CER and CO2
concentration in the leaf chamber, except for cassava.
Thus, it appears that factors other than CO2 level in
the leaf chamber cause negative correlations between
LA and CER.
Leaf area also showed negative correlations with
dark respiration rates for wheat (22), alfalfa (10), and
peanut (7). These negative correlations also indicate
that metabolic differences are responsible for the LA-
CER relationships since CO2 concentration changes
associated with differences in LA and constant flow
rates would not influence dark respiration.
Transpiration and LA were inversely related in
soybean (5) (r = —0.94), but not in wheat (r = —0.05)
(22). Negative correlations of LA with leaf conduct-
ance (stomatal and boundary layer) of soybean (5)
indicate that large leaves have more resistance to CO2
movement through stomata than small leaves. The
apparent control of stomatal opening by reactions
related to photosynthetic capacity of the mesophyll
(31) may mean that the relationship of LA and leaf
conductance is indirect and mediated by photosyn-
thetic metabolism.
Most of the data used in this report were from
genotype comparisons in which the youngest fully-
expanded leaves near stem apices were measured.
Since leaf position on the stem often greatly influ-
ences leaf size and the degree of mutual shading, a
bias is possible which results from larger leaves being
at lower stem positions and thus, older, more heavily
shaded, and lower in CER. This bias is unlikely, how-
ever, since leaf selections for genotype comparisons
are carefully made to represent similar age and stem
position. It is especially unlikely in comparisons in-
volving flag leaves of single plants of cereal grains (2,
3) in which mutual shading would be nearly non-
existent. In addition, some experiments in which the
negative correlation between LA and CER were ob-
served were with plants young enough that successive
leaves were likely to be larger, not smaller (6).
The effects of LA on photosynthesis and crop pro-
ductivity needs further research. Growth analysis (20)
and modeling studies (9) for cotton indicated that the
okra leaf character (usually smaller leaves) has a de-
cided agronomic advantage over normal leaves, which
generally are larger. On the other hand, narrow and
broad leaflet isolines of soybean gave similar yield
(23), and in barley, no consistent difference in CER
and grain yield was observed between two groups of
lines selected for large and small leaves (3). Thus,
advantages of small or large leaves in crop production
are still open to question.
The negative correlations between CER and LA
occurred in almost all leaf photosynthesis studies sur-
veyed. Comparisons of CER involving genotypes with
different leaf sizes, may not indicate the inherent dif-
ferences in photosynthetic potential. The negative
correlations between LA and CER may be one of the
causes for lack of consistent correlations between
photosynthesis and yield. Thus, before undertaking
screening of crop germplasm for CER, preliminary
experiments should be performed to evaluate the
possible confounding effects of LA on CER.
132 CROP SCIENCE, VOL. 26, JANUARY-FEBRUARY 1986