24.04.2013 Views

A review of dipterocarps - Center for International Forestry Research

A review of dipterocarps - Center for International Forestry Research

A review of dipterocarps - Center for International Forestry Research

SHOW MORE
SHOW LESS

You also want an ePaper? Increase the reach of your titles

YUMPU automatically turns print PDFs into web optimized ePapers that Google loves.

Seed Physiology 61<br />

§ orthodox; and<br />

§ recalcitrant.<br />

The orthodox type is capable <strong>of</strong> desiccation to a low<br />

moisture content (approximately 5%) and storage <strong>for</strong><br />

several years at -20 o C with little loss <strong>of</strong> viability (Roberts<br />

1973). By contrast, the recalcitrant type is not capable<br />

<strong>of</strong> desiccation to a low moisture content without loss <strong>of</strong><br />

germination capacity and cannot be stored <strong>for</strong> long<br />

periods <strong>of</strong> time (Roberts 1973).<br />

A third category <strong>of</strong> seed storage physiology has been<br />

described. It was first defined in 1984 in relation to<br />

Araucaria columnaris seed (Tompsett 1984) and was<br />

termed ‘orthodox with limited desiccation ability’<br />

(OLDA). A similar category was later defined <strong>for</strong> c<strong>of</strong>fee<br />

seed and termed ‘intermediate’; the name denotes its<br />

partial tolerance <strong>of</strong> desiccation (Ellis et al. 1990, 1991).<br />

Some recent evidence (Tompsett, unpublished), however,<br />

confirms there may be little physiological difference<br />

between this third category <strong>of</strong> seed and the orthodox<br />

type. There are, however, important practical handling<br />

difficulties associated with this third category. These<br />

problems justify its retention as a distinct storage type.<br />

Some tropical seed is additionally subject to lowtemperature<br />

damage when stored in the moist condition<br />

(chilling damage, see pages 58-59). As a result, further<br />

categories could have been included. However, it was<br />

considered preferable, <strong>for</strong> the sake <strong>of</strong> simplicity, to<br />

employ only the three desiccation-damage-based<br />

categories described above. To date, all dipterocarp<br />

species examined have been found to be subject to<br />

chilling damage when moist.<br />

Desiccation physiology<br />

Curves <strong>of</strong> germination percentage against moisture<br />

content percentage can be plotted <strong>for</strong> the results from<br />

controlled desiccation studies. These curves illustrate<br />

whether the seed is recalcitrant or not and give<br />

parameters <strong>for</strong> the way the seed responds when it is dried.<br />

One parameter is the lowest-safe moisture content<br />

(LSMC), defined as the value below which viability is<br />

immediately lost on drying. The value <strong>for</strong> this parameter<br />

provides a guide to the moisture content below which<br />

seed should not be held during handling procedures.<br />

LSMC values were assessed under standard drying<br />

conditions and were found to vary between 26% and 50%<br />

(Table 3). In Table 4 further LSMC data are given;<br />

although these were assessed using various desiccation<br />

methods, the results are in broad agreement with those<br />

in Table 3.<br />

Slope and intercept parameters are presented <strong>for</strong><br />

some species (Table 3); these define the relationship<br />

between germination and moisture content during<br />

desiccation.<br />

Desiccation rates<br />

It is possible that desiccation rate may influence viability;<br />

<strong>for</strong> example, seeds dried quickly might give lower<br />

germination than seeds dried more slowly and gently to<br />

the same moisture content. However, in the case <strong>of</strong> the<br />

‘recalcitrant’ seed <strong>of</strong> Araucaria hunsteinii<br />

(Araucariaceae) no such differences were observed<br />

(Tompsett 1982). No intensive study <strong>of</strong> this sort has been<br />

carried out on dipterocarp seed. However,<br />

Amata-Archachai and Hellum (unpublished) found that<br />

immature fruits <strong>of</strong> Dipterocarpus alatus clearly dried<br />

quicker than mature fruits; they suggested that the<br />

difference could be because <strong>of</strong> the death on drying <strong>of</strong><br />

the immature seeds. However, the faster loss <strong>of</strong> moisture<br />

by immature seeds could also be explained by their<br />

smaller size. Small seeds have a higher ratio <strong>of</strong> surface<br />

area to volume than large seeds, enabling quicker<br />

moisture loss. In this connection, Tamari (1976) found<br />

small seeds <strong>of</strong> S. parvifolia (0.3 g) gave low viability,<br />

whilst large seeds (0.5 g) gave higher viability. One<br />

explanation <strong>for</strong> the latter finding is that the smaller seeds<br />

had dried quicker and thus lost more viability than larger<br />

seeds prior to testing.<br />

Clear-cut differences in desiccation rates among<br />

seeds <strong>of</strong> species in the same genus have been reported<br />

by Tompsett (1986, 1987); rates <strong>for</strong> Dipterocarpus<br />

seeds varied greatly and depended on their size and<br />

structure. At the one extreme D. intricatus seed required<br />

only one week to dry to 7% moisture content; at the other<br />

extreme, seed <strong>of</strong> D. obtusifolius under identical<br />

conditions retained c. 30% moisture content even after<br />

5 weeks. Likewise, Yap (1986) found S. parvifolia seeds<br />

dried quicker than those <strong>of</strong> two larger-seeded species<br />

<strong>of</strong> Shorea; he believed the difference in rates to be<br />

related to pericarp thickness.<br />

Differences in desiccation rates <strong>of</strong> the type discussed<br />

above may possibly affect both the initial<br />

post-desiccation viability and the subsequent storage life<br />

<strong>of</strong> the seed. Further studies are needed to assess these<br />

effects.

Hooray! Your file is uploaded and ready to be published.

Saved successfully!

Ooh no, something went wrong!