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Yoshida - 1981 - Fundamentals of Rice Crop Science

Yoshida - 1981 - Fundamentals of Rice Crop Science

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234 FUNDAMENTALS OF RICE CROP SCIENCE<br />

Table 7.1. Photosynthetic activity <strong>of</strong> leaves, sheaths, and panicles at a light intensity<br />

<strong>of</strong> 60 klx at milky stage. a<br />

Leaf blade 1 c<br />

2<br />

3<br />

4<br />

All leaves<br />

Leaf sheath plus culm<br />

Panicle<br />

Total<br />

Surface<br />

area<br />

(cm 2 )<br />

24<br />

54<br />

51<br />

37<br />

Assimilation b<br />

Rate<br />

(mg CO 2 /h) (mg CO 2 /dm 2 per h)<br />

P n R d P g P n R d<br />

8.3 (30) 0.44 (9) 8.7 (27) 34 1.8<br />

10.1 (36)0.52 (11) 10.6 (33) 19 1.0<br />

5.6 (20)0.46 (11) 6.1 (19) 11 0.9<br />

2.2 (8) 0.34 (7) 2.5 (8) 6 0.9<br />

26.2 (94) 1.76 (38) 27.9 (87)<br />

1.2 (4) 0.60 (14) 1.8 (5) – –<br />

0.6 (2) 2.05 (48) 2.7 (8) – –<br />

28.0(100) 4.46(100) 32.4(100) – –<br />

a <strong>Yoshida</strong> and Cock (1971); variety IR22 grown in the field. b P n = net photosynthesis, R d =<br />

dark respiration, P g = gross photosynthesis. Figure in parentheses indicate percentage<br />

<strong>of</strong> the total. c Counted from the top.<br />

Table 7.1). The potential net photosynthesis <strong>of</strong> the leaves was 94% <strong>of</strong> the total.<br />

The fourth leaf had a very low net photosynthesis both on a per-leaf basis and on a<br />

per-unit-area basis. The top leaf had the highest net photosynthesis per unit area<br />

but the second leaf was larger in size and had a greater potential net photosynthesis<br />

on a per-leaf basis. In rice, all the leaves from the flag leaf down to the third leaf<br />

from the top export assimilates to the panicle. Lower leaves send their assimilates<br />

to the roots (Tanaka 1958). The top 3 leaves <strong>of</strong> plants in an IR8 crop made up<br />

74% <strong>of</strong> the total leaf area when LAI was 5.5 at heading. Flag leaves were only 19%<br />

<strong>of</strong> the total leaf area, and the second and third leaves made up 55% (<strong>Yoshida</strong> and<br />

Cock 1971).<br />

The net photosynthesis <strong>of</strong> the leaf sheath and panicle is extremely small. Even in<br />

terms <strong>of</strong> gross photosynthesis ( P g ), the panicle (8%) and the leaf sheath (5%) have<br />

extremely low photosynthetic capacity. Hence, both can be considered nonphotosynthetic<br />

tissues (Tsuno et al 1975).<br />

The longevity <strong>of</strong> the green tissue must be considered when assessing the<br />

photosynthetic contribution <strong>of</strong> different plant parts during ripening. The panicle<br />

becomes yellow relatively early in ripening but the leaves remain green much<br />

longer. The flag leaf remains green and maintains active photosynthesis until<br />

maturity (Takeda and Maruta 1956). At later stages <strong>of</strong> ripening, however, photosynthesis<br />

may not make much <strong>of</strong> a net contribution to grain production because<br />

rapid grain growth occurs earlier and slows down toward maturity.<br />

The light environment <strong>of</strong> different plant parts in a crop canopy is extremely<br />

important for determining the real photosynthetic activity <strong>of</strong> a given part. Panicles<br />

<strong>of</strong> improved indica varieties tend to bend and become positioned below the flag<br />

leaf where they are heavily shaded by the leaf canopy, particularly when the LAI is<br />

large. Panicles <strong>of</strong> these varieties are unable to make a significant contribution to<br />

grain filling because <strong>of</strong> their low potential photosynthetic activity and low light

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