Limiting Factors in Photosynthesis: California
Limiting Factors in Photosynthesis: California
Limiting Factors in Photosynthesis: California
NORMAN TERRY
Department of Soils and Plant Nutrition, University of California, Berkeley, Calf'ornia 94720
o0 0 0~~
0
0
~~~0 varied with the Chl content of the leaf: leaves with smaller ChM
m contents tolerated less irradiance than those with higher Chl
+ 0
30 F- 00 contents (Fig. 3). P. data were, therefore, obtained at irradiances
-J 00 0 appropriate to the Chl content of each leaf. The high ambient CO2
-J 0 00 concentrations used in the photosynthesis measurements (1,000
a. 00 6 ,ul CO2 I1 air) ensured that rates of photorespiration were minimal.
0 20k The data show that Pm./area is linearly related to Chl/area
a:
I 0~~~~
(Fig. 4A). The linear regression of the data is significant to P =
) 0b00 0.001 with a correlation coefficient of 0.95. For ease of discussion
1o0 e 00 the relationship between P./area and Chl/area is referred to as
"lineai" but it should be pointed out that it is almost certainly
slightly curvilinear. Statistically, the fit to the data was improved
s 0 I I
significantly by the addition of a quadratic term to the regression.
0
1.0 2.0 3.0 4.0 Furthermore, the quadratic regression (y = -4.93 + 2.57 x -
0.014 x2 yielded a negative rather than a positive intercept; a
LEAF Fe CONCENTRATION(pmol g-1 dry wt) negative intercept is more appropriate since P would attain a
FIG. 1. Relation between Chl content and leaf blade Fe concentration negative value (i.e. respiration) when leaves reached very low Chl
(y = 2.29 X3 - 20.1 X2 + 63.4 x - 16.7). contents (<2 lag cm-2).
Although Fe stress decreased the rate of photosynthesis per unit
area, it did not decrease the rate of photosynthesis per unit Chl
0
A. (Fig. 4B). The rate of photosynthetic CO2 fixation remained high
at 437 98 ,umol CO2 mg Chl-M h-1 throughout the range of leaf
N 4.01- Chl contents. These data suggest that the photosynthetic apparatus
Eu associated with Chl functioned efficiently even at the very low leaf
C3% Chl contents associated with severe Fe stress.
I 3.01- 0 Leaf and Chloroplast Attributes. There are several possible
-J ways in which P,../area might be causally related to Chl/area.
0i 0 First, Fe stress might reduce the number or size of cells resulting
I 0
in less leaf tissue per unit area, thereby decreasing both Chl and
2.0h
0 0 0 photosynthetic capacity per unit leaf area. Second, Fe stress might
reduce the number or amount of chloroplasts per unit volume leaf
z 1.0 H tissue. A third possibility is that Fe stress might diminish the
x c amount of light-harvesting and electron transport apparatus (i.e.
0 I l Il I -
B. 400
E
N
100C) 80
N 4.0 t-
E u 60
E
Lu 3.0 Un
U) 40
w
z 0
I
o 2.0 0
0
X-
z 20
0
ci: 0
U)
0 4.0 e
0
0 0
0
I
a0 0A -20
L-
oII I I 0 4000 2000 3000 4000 5000
o0o 10 20 30 40 50 60 INCIDENT QUANTUM FLUX DENSITY
CHLOROPHYLL (A+B) CONTENT (p.E m-2s-4)
(.Lg cm-2) FIG. 3. Relation between the rate of photosynthesis and irradiance for
FIG. 2. Relationship between carotenoid pigment content and Chl a high Chi (50 yg cm-) control leaf and for a low Chl (6.4 jtg cm-) Fe-
content under conditions of Fe stress. A: xanthophyll (y = 1.27 + 0.0436 stressed leaf. Photosynthesis was measured as leaf CO2 uptake at 30 C at
x; r = 0.85); B: carotene (y = 0.182 + 0.0508 x; r = 0.94). saturating levels of CO2 supply (1,000 ,ul CO2 1` air).
Downloaded from on September 30, 2017 - Published by www.plantphysiol.org
Copyright 1980 American Society of Plant Biologists. All rights reserved.
Plant Physiol. Vol. 65, 1980 LIMITING FACTORS IN PHOTOSYNTHESIS 117
Table I. Effect of Fe Stress on Some Leaf and Chloroplast Attributes
120 A
Control Fe-stressed Severely Fe-
0 Leaves Leaves stressed
>40 jig Chl 20-40ug Chl <20 ug Chl
1001- 0 cm-2 cm-2 cm-2
WI 0
N= 14 N= 12 N=8
X I~N
80s- No. of cells/leaf area (106 1.30 +0.30 1.21 0.19 1.35 0.28
.-- 0 C cm2)
l IE
.4Q 00
0 9 00 Mean leaf cell volume (10-8 2.64 0.75 2.78 0.61 2.75 0.90
00 cm3)
601-
No. of chloroplasts/leaf area 1.02 0.19 0.91 0.21 0.89 0.18
0~~~~~ (108 cm-)
00 No. of chloroplasts/cell 72 32 77 9 83 24
401-
Average chloroplast volume 42 8.3 37 12 21 7.3
0 (A)
o&
20 Soluble protein/leaf area (mg 0.57 0.08 0.56 0.07 0.53 0.07
cm-)
N=6 N=6 N=7
0
I l I Nonaqueously isolated:
B Dry wt/chloroplast (pg) 30.1 5.2 23.0 4.8 17.4 8.1
4000
E Protein N/chloroplast (pg) 1.88 0.37 1.34 0.25 1.24 0.46
E 800 0
_ 600
x 400 0.8
4 _ 0-o 0
0- o 200
U- 0
l I I I I
10 20 30 40 50 60 0.61 o
CHLOROPHYLL (A+B) CONTENT (pg cm-2) I
-J
0
FIG. 4. Relationship between maximum photosynthesis and Chl con- 0
tent under conditions of Fe stress. A: rate expressed per unit leaf area (y 0
0.
cx 0.4
= 4.97 + 1.69 x; r = 0.95); B: rate expressed per unit Chl (y = 482 - 1.53 -J,
x; r = -0.26). C-)
0~~~~~
0.21
membranes, pigments, reaction centers and electron carriers, etc.)
per chloroplast. The latter possibility might result in a reduction 0 00
0
in light absorption, or, photochemical capacity. These possibilities
0~~~~~~
were explored experimentally.
The data show that Fe stress did not affect the number of cells/ 2.4
leaf area or average leaf cell volume, nor did it change the number 0 0~~~~~~ 0
of chloroplasts/area or per cell (Table I). In an earlier experiment
it was observed that for sugar beet leaves of comparable size and 4.6 [ _0 00
l
C~~~~~~
_
development, Fe stress had no effect on fresh weight/area, leaf 0
thickness, or leaf tissue volume, i.e. volume exclusive of leaf Q-j
0~~~~~~
intercellular air space (24). Collectively, these data imply that Fe ~-0
0.8
stress did not diminish Chl/area and photosynthetic capacity via
an effect on the amount of leaf tissue, or the number of chloro-
plasts per unit area. cr-0~~~~~
0~~~~~~~
There was some evidence that Fe stress reduced the size of 24 00 0O0B0
chloroplasts. Direct measurement of the radii of the individual 8 O
chloroplasts yielded an estimated volume which was smaller for 00 00
severely Fe-stressed than for mildly stressed or control leaves 161 ~jN 00
(Table I). In order to obtain an alternative measure of chloroplast 0 0,1 01 1 l
size, chloroplasts were isolated nonaqueously and the dry weight
per chloroplast determined; these data show a progressive decrease 8
XC-)
The reduction in Chl/area was due to a decrease in Chl/ + 0.01 x; r = 0.88); B: amount of protein N per chloroplast (y 0.99 + =
chloroplast (Fig. 5A). Thus, the relation of P,/area with Chl/ 0.016 x; r = 0.65); C: per unit leaf area rate of CO2 fixation by extracted
area is more specifically a relation between P../area and Chl/ RuBP carboxylase (y = 12.5 + 0.13 x; r = 0.65).
Downloaded from on September 30, 2017 - Published by www.plantphysiol.org
Copyright 1980 American Society of Plant Biologists. All rights reserved.
118 TERRY Plant Physiol. Vol. 65, 1980
4.0
0.8
673~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~T
664 ~~~~~~~~~E
z~~~~~~~~~
LX
0.2 630
m522 ~~~~~~~~~~~~~~~LLJ
0L 10 20 30 4 0 6
0 IZ
0 10 20 30 40 50 60