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Photosynthesis pp 301–320Cite as

Fructans: Synthesis and Regulation

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Part of the book series: Advances in Photosynthesis and Respiration ((AIPH,volume 9))

Summary

Roughly ten percent of higher plant species possess a mechanism for reserve carbon allocation based on soluble fructose polymers (vacuolar fructan), which differs markedly in its enzymology, subcellular compartmentation and regulation from the more common starch-based carbon economy. This includes many economically important species, especially the temperate grasses and cereals. This chapter describes these novel elements associated with fructan metabolism, with particular emphasis on fructan synthesis in photosynthetic tissue.

Fructan structures, although based on variations in a few characters (polymer size, glycosidic linkage, etc.) are shown to be both varied and complex, differing markedly between species but possessing a consistency within species which argues for a biosynthetic mechanism with a high degree of specificity. The enzymological mechanisms currently thought to be involved are discussed, with particular reference to the involvement of multifunctional enzymes and the strong effects of both substrate and enzyme concentration on the chemical nature of the products in vitro. The enzymatic polymerization of authentic grass fructan has been achieved, but the conditions required in vitro do not coincide with those expected in the vacuole, the currently accepted site of synthesis. The properties of fructosyl transferases in general are shown to be unusual and we emphasize the need toreconcile the characteristics of enzymes in vitro with the patterns of metabolism and conditions observed in the tissue.

The regulation of fructan synthesis is discussed in relation to the pivotal role of sucrose as the sole substrate and as the key element in the coarse control of fructan accumulation, apparently acting at the level of gene expression and de novo enzyme synthesis. Sucrose mediated feedback inhibition of starch metabolism via phosphate transport does not apparently occur in grass leaves. This isolation of chloroplast metabolism from cytosolic sucrose accumulation indicates a fundamental difference in the fine control of centralcarbon metabolism between fructan and starch accumulators.

Recently, non-fructan, starch-accumulating plants such as maize, spinach and tobacco have been transformed with bacterial genes for fructan synthesis and shown to accumulate fructan. The current value of such transgenics is in terms of what they may tell us about the regulation of primary carbon metabolism in the recipient plants. Transgenics currently provide little insight into the nature or control of endogenous fructan metabolism. Some neglected aspects of the physiology these transgenics are considered by comparison with endogenous reserve carbon metabolism in untransformed plants.

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Abbreviations

DP:

degree of polymerization (number of hexose moieties)

FFT:

fructan:fructan fructosyl transferase (EC 2.4.1.99)

Mr:

molecualr mass

PEG:

polyethylene glycol

SST:

sucrose:sucrose fructosyl transferase (EC 2.4.1.100)

TLC:

thin layer chromatography

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Authors and Affiliations

  1. Environmental Biology Department, Institute of Grassland and Environmental Research, Plas Gogerddan, Aberystwyth, SY23 3EB, Wales, UK

    A. J. Cairns, C. J. Pollock, J. A. Gallagher & J. Harrison

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  1. A. J. Cairns
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  2. C. J. Pollock
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  3. J. A. Gallagher
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  4. J. Harrison
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Editors and Affiliations

  1. Robert Hill Institute, University of Sheffield, UK

    Richard C. Leegood

  2. Department of Botany, University of Wisconsin, USA

    Thomas D. Sharkey

  3. Research School of Biological Sciences, Australian National University, Australia

    Susanne von Caemmerer

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Cairns, A.J., Pollock, C.J., Gallagher, J.A., Harrison, J. (2000). Fructans: Synthesis and Regulation. In: Leegood, R.C., Sharkey, T.D., von Caemmerer, S. (eds) Photosynthesis. Advances in Photosynthesis and Respiration, vol 9. Springer, Dordrecht. https://doi.org/10.1007/0-306-48137-5_13

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