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The polysaccharide storage form of glucose in animals is glycogen, whereas in plants it is starch. Both of these are polymers of α-glucose with α-l,4 glycosidic linkages and α-l,6 glycosidic branch points (Wikipedia article on polysaccharides). The only difference that most sources mention (e.g. Berg et al.) is that glycogen contains more branches than starch.

It is not clear to me from this information what effect the different branching would have on the structures of the polysaccharides, nor why one rather than the other would be preferred in animals and plants.

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  • $\begingroup$ It is surprisingly difficult to find a proper answer to this question on the internet — my own answer was only found after consulting specialized reviews. I therefore think this is an important question and have therefore edited it, tightening up the wording, avoiding the implication that polysaccharides are the only storage form, and spelling out the chemistry. If the original poster is still active on the list I hope he will accept these changes. $\endgroup$ – David Jan 9 '18 at 17:30
  • $\begingroup$ What about fungi? Do they even have storage polysaccharides and if so, what kind? $\endgroup$ – jaia Jan 12 '18 at 2:03
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Summary

The key difference between glycogen and amylopectin (the main constituent of starch) is not the number of α l,6-glycosidic branches, but their arrangement.

In glycogen branches are successively subdivided, producing a relatively small globular structure that is unable to grow further. It is soluble in an aqueous environment and, with its numerous exposed ends, can be metabolized rapidly — appropriate for animal cells in which energy reserves must be mobilized in response to immediate demands, e.g. for muscle contraction.

In amylopectin there is a long central polysaccharide chain from which branches of limited size extend at intervals. This produces much larger semi-crystalline particles (starch grains), a form especially suited to long-term bulk storage in seeds and tubers.

The Chemistry

Chemistry of glycogen and starch

This is the common feature of glycogen and the amylopectin portion of starch. (The amylose portion is unbranched.) In glycogen there is approx. one branch point per 10 glucose units, whereas in amylopectin the figure is 1 per 24–30 (source: Wikipedia).

The Topography

The contrasting branching topography of the two polysaccharides, mentioned above, is illustrated diagrammatically below:

Topography of branching in glycogen and starch

This is a two-dimensional representation. In three dimensions the glycogen spreads out in all directions from a central point — actually the primer enzyme, glycogenin. In three-dimensions the amylopectin strands mainly lay side by side.

The Macro-structure

The illustration below, modified from Bell et al., shows the different shapes and sizes of the macromolecular structures. It should be mentioned that semi-crystalline nature of amylopectin is aided by the helical conformation of the chains.

Macrostructure of glycogen and starch

Rather than providing a précis of the review of Bell et al. (Journal of Experimental Botany, Vol. 62, pp. 1775–1801, 2011) I shall quote from them directly (omitting their citations).

As regards glycogen they write:

Each chain, with the exception of the outer unbranched chains, supports two branches. This branching pattern allows for spherical growth of the particle generating tiers (a tier corresponds to the spherical space separating two consecutive branches from all chains located at similar distance from the center of the particle). This type of growth leads to an increase in the density of chains in each tier leading to a progressively more crowded structure towards the periphery.

Mathematical modelling predicts a maximal value for the particle size above which further growth is impossible as there would not be sufficient space for interaction of the chains with the catalytic sites of glycogen metabolism enzymes. This generates a particle consisting of 12 tiers corresponding to a 42 nm maximal diameter including 55,000 glucose residues. 36% of this total number rests in the outer (unbranched) shell and is thus readily accessible to glycogen catabolism without debranching. In vivo, glycogen particles are thus present in the form of these limit size granules (macroglycogen) and also smaller granules representing intermediate states of glycogen biosynthesis and degradation (proglycogen). Glycogen particles are entirely hydrosoluble and, therefore, define a state where the glucose is rendered less active osmotically yet readily accessible to rapid mobilization through the enzymes of glycogen catabolism as if it were in the soluble phase.

Regarding amylopectin they write:

Amylopectin defines one of, if not the largest, biological polymer known and contains from 105–106 glucose residues. There is no theoretical upper limit to the size reached by individual amylopectin molecules. This is not due to the slightly lesser degree of overall branching of the molecule when compared to glycogen. Rather it is due to the way the branches distribute within the structure. The branches are concentrated in sections of the amylopectin molecule leading to clusters of chains that allow for indefinite growth of the polysaccharide. Another major feature of the amylopectin cluster structure consists of the dense packing of chains generated at the root of the clusters where the density of branches locally reaches or exceeds that of glycogen. This dense packing of branches generates tightly packed glucan chains that are close enough to align and form parallel double helical structures. The helices within a single cluster and neighbouring clusters align and form sections of crystalline structures separated by sections of amorphous material (containing the branches) thereby generating the semi-crystalline nature of amylopectin and of the ensuing starch granule. Indeed the crystallized chains become insoluble and typically collapse into a macrogranular solid. This osmotically inert starch granule allows for the storage of unlimited amounts of glucose that become metabolically unavailable. Indeed the enzymes of starch synthesis and mobilization are unable to interact directly with the solid structure with the noticeable exception of granule-bound starch synthase the sole enzyme required for amylose synthesis.

Coda

The paucity of information on plant starch metabolism would seem to reflect a combination of their being less is known about plant biochemistry, and less general interest because of a general focus on medical and animal biochemistry. Although an animal biochemist myself (and, thus, previously ignorant of the information in this answer) I feel that it is time to redress this imbalance.

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  • $\begingroup$ Your labels for the alpha 1>4 and alpha 1>6 linkages in the first diagram are the wrong way around. $\endgroup$ – Alan Boyd Feb 7 '18 at 16:00
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well glycogen can be broken down into sugars a lot faster, many more branches means many more ends to clip individual sugars off of, that's how you mobilize the sugar for use, it is clipped of the end of a strand. With many more branches glycogen can mobilize more sugar more quickly. This is not important in plants but in animals that need to be able mobilize lots of energy in a hurry, glycogen works better. Additionally glycogen is a smaller molecule and easier to make, not surprising since glycogen is the ancestral condition for plants and animals.

As for why plants switched to starch, or more precisely gained it through symbiosis, starches folded crystalline structure makes it a higher density energy store but also slows its release, it is however more stable, which is important if you are going to be storing it for a long time. Animals would likely switch to starch too if they did not have to break it down to digest it, just because they take in so much of it. But since they have to break it down there is no real incentive the build it back into starch when glycogen has some advantages and quite frankly since the pathway is already there and evolution has a lot of "eh good enough".

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  • $\begingroup$ Any references for either of the claims? $\endgroup$ – another 'Homo sapien' Oct 1 '17 at 12:01
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    $\begingroup$ Added, and good thing to my old notes were wrong about ancestral condition. $\endgroup$ – John Oct 1 '17 at 15:40
  • $\begingroup$ Some interesting ideas, but seems speculative. (1) It's true that glycogen can generally be mobilized faster than starch, but I'm not sure it's due to higher branching -- I would rather think it's due to the soluble nature of glycogen granules vs. the crystalline form of starch, which you mention later. (2) What exactly do you mean by glycogen being "easier to make"? (3) You seem to contradict yourself in saying that for animals "glycogen works better", and then that animals "would switch to starch too" if they could? $\endgroup$ – Roland Dec 1 '17 at 16:35
  • $\begingroup$ Sorry, (1) I meant evolutionarily speaking animals probably would just use starch as it is if they did not have to take it apart then rebuild it to get it inside, since the advantage of glycogen is nominal. (2) Glucose is a simpler molecule with a shorter synthesis pathway. One step in starch synthesis involves producing amylopectin a molecule very similar to glucose. (1)the branching and the solubility are related and both definitely contribute, but starch is also a more complex molecules which requires multiple enzymes and must be unwound to be broken down into glucose. $\endgroup$ – John Dec 1 '17 at 21:09
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    $\begingroup$ @David please read the full exchange you are basically restating what I already said. The reason they never evolved it is they gain nothing by rebuilding glucose into starch instead of glycogen, the bonds are identical but the finished molecule and there properties are not. $\endgroup$ – John Jan 9 '18 at 21:04

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