The role of carbohydrate storage and redistribution in the source‐sink relations of wheat and barley during grain filling — a review

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Abstract
SUMMARY: Two types of source contribute photosynthate for grain filling in wheat (Triticum aestivum L.) and barley (Hordeum vulgare L.), current photosynthate transferred directly to the grain and photosynthate redistributed from reserve pools in vegetative tissues. Reserve pools provide the substrate needed to maintain transport and the supply of assimilate to grains during the dark period of the diurnal cycle and during the latter part of grain filling, when the photosynthetic apparatus is senescing and the rate of dry matter accumulation of grains exceeds the rate of dry matter accumulation of the total crop. In addition reserve pools provide a means by which the current rates of photosynthate production and of photosynthate use in grain filling are allowed to proceed (at least in part) independently from each other. There is evidence that all photosynthetic organs (leaves, glumes and exposed portion of the peduncle) contain one or more diurnal carbohydrate storage pools. Diurnal storage of sucrose seems to be much more important than the transient storage of starch. There is little evidence for fructan pools serving a role as a net source of carbon during the dark period of the diurnal cycle. However, fructan is the most important longer‐term reserve carbohydrate of vegetative tissues. Fructan accumulation occurs mainly in the extended internodes and leaf sheaths and usually terminates within three weeks following anthesis, after which fructan is gradually lost until grain maturity. The accumulation and loss of fructan are greatly influenced by environmental conditions and treatments that alter the longer‐term balance between photosynthate production by the plant and photosynthate use in grain filling. Observations are consistent with the view that fructan accumulation in expanded vegetative tissue is not competitive with grain filling, but the fructan pools accept surplus photosynthate during periods of low demand by grains and provide photosynthate during longer‐term deficits in current photosynthate production. Even under optimal conditions for photosynthesis it is likely that half or more of the photosynthate in mature grains is temporarily deposited in one or more reserve pools before being transferred to the grain. Neither the efficiency of reserve utilization in grain filling nor its potential variability in different genotypes and environments are known. Also, very little is known about the mechanisms that control the partitioning of photosynthate between the grain and reserve pools and the allocation to different types of reserve pools. Future progress in the understanding of photosynthesis‐yield relationships will likely depend to a great extent on improved knowledge of the controls that govern photosynthate deposition and redistribution in the different pools of reserve carbohydrate.