Lecture 24

Physiology of riceyield determinant

Introduction
Rice is the staple food grain for more than half of the world's population. Rice is grown on 150 million hectares, more than 10% of the earth's arable land Rice accounts for about 30% of total cereal production, almost equal to the level of wheat production (FAO, 1994). Unlike wheat. 95% of the world's rice is grown in less developed nations, primarily in Asia. Developed countries produce only about 5% of the total yield in 3% of the total harvested area (IRRI, 1995).

 There are two species of cultivated rice, Oryza sativa and Oryza glaberrima (Morishita, 1984). O. sativa is the main species of rice cultivated widely throughout the world. O. glaberrima is endemic only to West Africa. These two species are classified into the AA genome line, but are subdivided phyletically into different series. O. sativa is understood to have originated from the wild species O. perennis, and O. glaberrima is thought to have arisen from the wild species O. breviligulatu.

 O. sativa is classified into the three basic types: japonica, indica, and javanica (Takahashi, 1984). classifications are based on geographical distribution and morphological traits. Japonica is mainly cultivated in temperate areas. Indica is mainly distributed in the subtropics and tropics, but recent progress in conventional breeding of rice varieties has resulted in a wider and more complex distribution of indica varieties. Recent evidence shows that the javanica rice group is a subgroup of japonica germplasm (Glaszman, 1987). Thus, the javanicas are often called "tropical japonicas".

Growth and development

3-6 months from germination to maturity, depending on the variety and environment. life history of rice has three agronomic growth stages: vegetative, reproductive, and grainfilling.

 vegetative stage refers to the period from germination to the initiation of panicle primordia; the reproductive stage, from panicle primordia initiation to heading; and the grainfilling stage, from heading to maturity. The potential size of crop yield is primarily determined in the reproductive stage. Actual yield is largely determined during the grain filling period.

 The vegetative stage is characterized by active tillering.

In transplanted rice, about 10-30 tillers may be produced at conventional spacing, and the point of maximum tiller number typically occurs just before or during panicle initiation stage.

 Differences in growth duration are primarily due to differences in the length of the vegetative growth stage.

 The reproductive stage is characterized by culm elongation, a decrease in tiller number, emergence of flag leaves (the last leaf), booting, heading, and flowering. Panicle development and growth start with initiation of panicle primordia and end when the pollen is fully matured. Initiation of panicle primordia usually occurs about 30 days before heading. Spikelet number, a yielddetermining factor, is determined during this period.

 The grainfilling stage is characterized by grainfilling and leaf senescence. The length of grainfilling, largely affected by temperature, ranges from about 30 days in the tropics to 65 days in cool, temperate regions. The proportion of filled spikelets is mainly determined during this period

Grain filling: concurrent photosynthesis and remobilization of pre-stored carbon

Grain filling is the final stage of growth in cereals where fertilized ovaries develop into caryopses. At this stage, about 40--50% of total biomass is formed. Grain filling is contributed by both the concurrent photosynthesis during this period and remobilization of pre-stored carbon in sheath and stems.

 Unlike other major cereal crops, rice has two hard and interlocked hulls that limit grain size; in wheat, grain size is much more variable and depends largely on the length of grain-filling period.

 Monocarpic plants, such as rice and wheat, need the initiation of whole plant senescence so that stored carbohydrates in stems and leaf sheaths can be remobilized and transferred to their grains.

 Normally in these crops, pre-stored food contributes 25-33% to the final weight of a grain. Delayed whole plant senescence, leading to poorly filled grains and unused carbohydrate in straws, is a new problem increasingly recognized in rice and wheat production in recent years.

 Slow grain filling is always associated with delayed whole plant senescence. Although farmers can choose cultivars of early-maturation, there are still some situations that have made the delayed senescence a serious problem that needs attention

1.It is well known that heavy use of nitrogen fertilizers can lead to delayed senescence and, in worst cases, canopy lodging. Although farmers are generally aware of this, the problem still occurs every year and everywhere, especially in highly productive areas. This is possibly related to the intensive nature of agriculture today: using less arable land to feed ever more people

2.Selection of lodging-resistant cultivars to cope with the lodging problem has led to another problem in some cases, namely that stems are short and strong but stored carbohydrate is poorly used because the plants may stay `green' for too long, particularly in some cases with short-grain rice cultivars.

3. Introduction of hybrid rice has been a fantastic success in China. Utilization of heterosis - a hybrid between the two subspecies (or ecotypes) of rice, the Japonica and Indica rices - has, however, also met the problem of delayed senescence. Grains are poorly filled or un-filled kernels. Such hybrid genotypes seem too vigorous in terms of keeping `young'.

 We may define cases described above as unfavorably delayed senescence, which means that no gain is obtainable from the extended grain-filling period. We should however, distinguish this situation from that in favorable conditions, in which early senescence should be avoided because it reduces the photosynthesis during the grain-filling period and therefore reduces grain weight (Zhang et al., 1998).

 Delayed senescence in 2 and 3 above is basically a genetically controlled characteristic and there are no practical measures to deal with this problem. Situation I is managed in practice with so called `appropriate control of nitrogen fertilizers and canopy density', which is often too late when crops are in the grainfilling stage.

 In all of these three situations, a slow grain filling is a yield-limiting factor and any way in which the rate of grain filling can be enhanced should be beneficial to the final yield formation

 If weather conditions are favorable, and if such delayed senescence is also `functionally' delayed, that is, photosynthesis and phloem translocation are functional, such delayed senescence may help achieve higher dry mass production and possibly higher grain yield.

 However, in many cases, kernels and their connecting rachis or rachilla seem mature or senesce earlier than the stem and leaves. Much unused carbohydrate is left in the stem and sheath.

Yield components
A useful method for examining yield performance is to break the yield into its components (Murata and Matsushima, 1975; Yoshida 1981 a) : Grain yield (t ha-') =

=panicle number m-2 x spikelet number/panicle x % filled spikelets x 1,000-grain weight(g) x 10-5 = spikelet number m-2- x% filled spikelets x I,000-grain weight (g) x 10-5

 Each yield component is largely established at a particular growth stage. transplanted rice, the number of panicles - is largely dependent on tiller performance. direct seeding system, the number of panicles per square meter is largely dependent on seeding rate and percent emergence.

 The number of spikelets per panicle, on the other hand, is determined during the reproductive growth stage. Early in the reproductive growth, the maximum number of spikelets is determined by the differentiation of branches and spikelets. after spikelet differentiation, some spikelets may degenerate. The number of spikelets observed at heading or at maturity is the difference between the number of differentiated primordia and the number of primordia that degenerate. The percentage of filled spikelets is determined in the period from anthesis to physiological maturity.

 Unfavorable weather conditions, such as low or high temperature at the reduction division stage and anthesis, may induce sterility. Unfavorable weather conditions or other stresses during grainfilling may reduce or terminate growth of some spikelets, resulting in unfilled spikelets.

 The 1,000-grain weight is a stable varietal character because the grain size is rigidly controlled by the size of the hull. Hence, grain cannot grow to a size greater than that

permitted by the hull

Harvest index and vield

The total dry weight of a rice crop under favorable conditions is around 10-20 t ha-', depending on variety, management, and environment. The harvest index is about 0.3 for traditional tall varieties and 0.5 for improved, semidwarf varieties. As a result, grain yield ranges between 3 and 10 t ha' per crop (Yoshida, 1981b).

 A grain yield of 16.6 t ha-' (rough rice) has been recorded with a japonica Fl hybrid rice in Yunnan Province, South China (2550'N, 10030'E, at 1640 m altitude) (Amano et al., 1996a, b). The harvest index was extremely high with a value of 0.67. In Australia and California, 13-15 t ha-' yield levels have been achieved (Kropff et al., 1994). The average rice yields of countries vary from 0.9 to 8.1 t ha-' (FAO, 1994) depending on climates, varieties, fertilizers and pesticide usage, other farming practices, etc.

Nitrogen nutrition and yield components

Increased rice production in the last 40 years is largely attributed to the increased use of N fertilizer. Nitrogen deficiency occurs almost everywhere unless N is applied as a fertilizer or manure. Rice plants quickly respond to N supply by change their pattern of growth. The amount of N uptake needed to produce one ton of rough rice is 15 to 17 kg N for an average yield 5-6 t ha -1 and 19 kg N

for high yielding rice (Wada et al., 1986).

 However, the absolute amount of N absorbed is not the only factor to be considered in high-yielding rice. Rather, the N supply pattern and uptake process of N throughout the whole life cycle of the plant is equally important. This is because formation (number, size, or extent) of each yield component strongly depends on the amount of N supply at each crucial stage for respective yield components.

There are two sources of N for field-grown rice plants, the soil and fertilizer. To obtain a high yield under given conditions, the supply of N from both sources should match the amount necessary to support the desired growth of the plant through all the stages of growth. During the last two decades, the absorption patterns of these two groups of N have been intensively studied throughout the life of the rice plant by using 15N as a tracer. It is now possible to describe how N from these two sources is absorbed by the plants, and its effects on the growth and yield components of irrigated rice grown in northeastern Japan (Mae and Shoji, 1984, Wada et al., 1986).

Relation between leaf photosynthesis and nitrogen Photosynthetic capacity per unit leaf area is thought to be an important factor related to crop productivity. The relationship between the rates of leaf photosynthesis and leaf N content has been repeatedly examined in rice leaves. Accumulated evidences indicate that the maximum rate of CO2 assimilation per unit leaf area (measured under light-saturated, ambient air conditions) is almost proportional or somewhat curvilinear to SLN (aount of N per unit leaf area (Specific leaf N)

 This must be due to the fact that a large part of leaf N is invested in photosynthetic apparatus, namely chloroplasts. In mature rice leaves, about 80% of the total leaf N is allocated to chloroplasts (Morita ,1980) as is the case with other C3 plants (Brown, 1978; Makino and Osmond, 1991).

Analysis of rate-limiting factors of CO, assimilation in rice leaves

Analysis of rate-limiting factors of leaf photosynthesis provides useful information for improving the productivity of rice plants. A biochemical model of C3 photosynthesis was proposed by Farquhar et al. (1980). According to their model, the rate of C02 assimilation under lightsaturated and low C02 concentration conditions is determined by the carboxylation capacity of ribulose1,5-bisphosphate (RuBP).

 This capacity is determined by the CO,_ diffusion process from the outside of a leaf to the carboxylation site of ribulose-1, 5bisphosphate carboxylase/oxygenase (Rubisco) in chloroplasts, and the amounts and kinetic parameters of Rubisco. Rubisco is a bifunctional enzyme which catalyze two enzyme competing reactions, photosynthesis and photorespiration. In addition to its important biochemical function, this enzyme is the most abundant protein in the leaves of C3 plants.

 The relationships between the rates of C02 assimilation, and the process of COz diffusion, Rubisco amounts and kinetic parameters were analyzed in rice leaves (Makino et al., 1985b), following the 0 photosynthesis model of Farquhar et al. (1980).

 The rates of COz assimilation under ambient COz concentration and light-saturated conditions could be predicted from the amounts and kinetic properties of Rubisco and leaf conductance. The results indicated that the amount of Rubisco is a rate-limiting factor of photosynthesis under ambient air and light-saturated conditions throughout the life cycle of the rice leaf.

 This conclusion was recently verified by gene manipulation experiments using an anti-sense gene for a small subunit of Rubisco in rice plants (Makino et al., 1997). The rate of COz assimilation under ambient air and light saturated conditions decreased with decreasing the amounts of Rubisco in the leaves of the Rubiscoreduced transgenic plants.

. The amount of Rubisco and its ratio to total

leaf' nitrogen in different environments

The SLN and N partitioning among photosynthetic components under different growth conditions are important factors related to photosynthetic N useefficiency (Evans, 1989; Hikosaka and Terashima, 1995).
The amount of leaf N and Rubisco per unit leaf area is affected by N supply. Greater N supply generally results in increased N and Rubisco per unit leaf area in developing leaves.

 Both the absolute amount of Rubisco and the proportion of Rubisco-N to total N in the fullyexpanded rice leaves increased from 22 to 32% with increased N supply during leaf growth. However, the N ratio for other photosynthetic components, such as chlorophyll and cytochrome f in proportion to total leaf N did not change.

 Under low irradiances, RuBP regeneration capacity, mediated by light harvesting and electron transport capacities, becomes a rate-limiting factor of leaf photosynthesis in C3 plants. Under elevated COz conditions, RuBP regeneration capacity, mediated by light harvesting, electron transport, and/or Pi regeneration capacities becomes a rate-limiting step (Farquhar et al., 1980; Sharkey, 1985). Under these conditions, Rubisco may exist in excess of the amount

 for RuBP regeneration capacity. If rice plants are potentially able to optimize N allocation under such different environmental conditions, N from excess Rubisco should be reallocated into limiting factors under respective conditions

Source and sink relationship

A temporary accumulation of starch and sucrose in the flag leaf was observed in panicle-free plants, but the accumulation was not significant. However, in the panicle-free plants, the weight of shoots, excluding panicles, increased by 200% and that of roots increased by 150%c, at the time of harvest.

 Thus, it appears that removal of panicle has no effect on the relationship between the rate of photosynthesis and the levels of the major biochemical participants in photosynthesis. These findings indicate that the sink capacity of panicles does not limit the source activity during grainfilling in rice plants.

 This leads us to the consideration that a lack of source capacity is the main constraint to increased rice yield potential when the grainfilling percentage of spikelets is not high in spite of favorable climatic conditions