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A review of dipterocarps - Center for International Forestry Research

A review of dipterocarps - Center for International Forestry Research

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Seed Physiology<br />

P.B. Tompsett<br />

Seed is the natural vehicle <strong>for</strong> gene movement and<br />

storage. It is the usual <strong>for</strong>m in which germplasm is<br />

collected. When procedures can be devised to transport<br />

and retain material in this <strong>for</strong>m, many <strong>of</strong> the technical<br />

problems associated with other methods can be avoided.<br />

This advantage renders seed especially appropriate <strong>for</strong><br />

users in tropical and subtropical countries. In general,<br />

seed is the most common <strong>for</strong>m <strong>of</strong> propagation <strong>for</strong><br />

af<strong>for</strong>estation and is the <strong>for</strong>m in which breeding stock is<br />

usually retained. There are, however, considerable<br />

problems remaining in the use <strong>of</strong> seed. Some <strong>of</strong> these<br />

are discussed below <strong>for</strong> dipterocarp species in relation<br />

to the underlying seed physiology processes.<br />

Much pioneering work on agricultural crop seed<br />

physiology was conducted over the last 20 years (see<br />

below <strong>for</strong> some references) and the principles<br />

discovered <strong>of</strong>ten apply to seed <strong>of</strong> woody species. These<br />

earlier results have been translated into technological<br />

principles. Thus, manuals have been published on the<br />

design <strong>of</strong> seed storage facilities (Cromarty et al. 1982),<br />

seed management techniques (Ellis et al. 1984) and a<br />

handbook on seed technology <strong>for</strong> genebanks (Ellis et al.<br />

1985). Knowledge <strong>of</strong> seed physiology has thus improved<br />

practical handling and management <strong>of</strong> crop seeds.<br />

Compared to crop species, relatively little research<br />

has been published on tropical and subtropical tree seed<br />

technology and physiology. Publications have been<br />

produced following IUFRO Seed Problems Group<br />

meetings, the most recent <strong>of</strong> which was held in Tanzania<br />

(Olsen 1996). Another source <strong>of</strong> in<strong>for</strong>mation is a<br />

summary <strong>of</strong> some relevant seed physiology projects<br />

which has recently been published in database <strong>for</strong>m<br />

(Tompsett and Kemp 1996a, b). Also, a seed compendium<br />

has been published which supplies succinct entries on<br />

many tropical trees (Hong et al. 1996).<br />

A considerable amount <strong>of</strong> empirical work on the<br />

storage <strong>of</strong> <strong>for</strong>est tree seed has been carried out; a<br />

sampling is given in Chapter 4. A more physiological<br />

research approach is relatively new. Many tree species<br />

have seed that is desiccation-sensitive (‘recalcitrant’),<br />

Chapter 3<br />

so that moisture physiology is especially important <strong>for</strong><br />

this group. However, in a recent <strong>review</strong> article on water<br />

in relation to seed storage, the section on desiccationsensitive<br />

seeds comprised only 4% <strong>of</strong> the article<br />

(Roberts and Ellis 1989). More attention is, however,<br />

now being given to recalcitrant seeds (see, <strong>for</strong> example,<br />

Berjak and Pammenter 1996).<br />

Framework <strong>of</strong> the Review<br />

Germination is basic to all aspects <strong>of</strong> seed studies; work<br />

on germination physiology, especially in relation to<br />

temperature, is thus considered first. Another important<br />

experimental consideration is the physiological<br />

condition <strong>of</strong> the seed at the time <strong>of</strong> harvest, moisture<br />

content being the single most important factor; this is<br />

considered next in the <strong>review</strong>. Thirdly, the effect <strong>of</strong><br />

desiccation is discussed; knowledge <strong>of</strong> this factor enables<br />

seed to be classed as orthodox (tolerant) or recalcitrant<br />

(intolerant). Finally, the effects <strong>of</strong> seed storage are<br />

considered.<br />

The Review<br />

Germination<br />

In general, dipterocarp seeds germinate quickly under<br />

moist, warm conditions.<br />

Early germination studies took the <strong>for</strong>m <strong>of</strong> nursery<br />

assessments leading to ecological conclusions. A major<br />

study <strong>of</strong> this type, in which 56 dipterocarp species were<br />

assessed <strong>for</strong> germination rate and final germination, is<br />

that <strong>of</strong> Ng (1980); no conclusions can be made<br />

concerning temperature effects. Conditions were more<br />

closely controlled in experiments on Shorea roxburghii,<br />

S. robusta and S. almon by Tompsett (1985); results<br />

showed optimum germination in the range 26-31°C <strong>for</strong><br />

these three species. Corbineau and Come (1986) found<br />

that final germination reached nearly 100% <strong>for</strong> both<br />

Hopea odorata and S. roxburghii over a broad range <strong>of</strong><br />

temperatures, but 30-35°C was deemed optimal because<br />

germination rates were faster. More recently, a standard

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