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A review of dipterocarps - Center for International Forestry Research

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Root Symbiosis and Nutrition<br />

in Sabah showed that increasing concentrations <strong>of</strong><br />

fertiliser resulted in increased growth rates but that<br />

growth was reduced when 2000 mg <strong>of</strong> NH 4 NO 3 was<br />

applied (Yap and Moura-Costa, in press). It will be<br />

interesting to see the final outcome <strong>of</strong> this large-scale<br />

field experiment.<br />

Lee and Lim (1989) found that foliar P concentration<br />

in naturally regenerated seedlings <strong>of</strong> Shorea curtisii and<br />

S. leprosula growing in a logged over <strong>for</strong>est site with<br />

low levels <strong>of</strong> available P (5.8 to 7.1 ppm) was significantly<br />

correlated with the extent <strong>of</strong> ectomycorrhizal infection.<br />

Lee and Alexander (1994) working with Hopea helferi<br />

and H. odorata found positive growth responses to<br />

mycorrhizal infection but variable responses to nutrient<br />

treatments. They also reported the first direct<br />

experimental evidence that ectomycorrhizal infection<br />

improved P uptake and growth <strong>of</strong> a dipterocarp species,<br />

H. odorata. Scleroderma dicstyosporum and S.<br />

columnare were reported to increase levels <strong>of</strong> nitrogen,<br />

phosphorus and potassium in seedlings <strong>of</strong> Shorea<br />

mecistopteryx (Supriyanto et al. 1993b) but these results<br />

may have been misinterpreted. Increased plant height<br />

growth, diameter and dry weight as well as uptake <strong>of</strong> Fe,<br />

Mn, Cu, Zn and Al by seedlings <strong>of</strong> Shorea compressa, S.<br />

pinanga, S. stenoptera, H. odorata and Vatica<br />

sumatrana inoculated with chopped fruit bodies <strong>of</strong><br />

Russula sp., Scleroderma sp. and Boletus sp. have been<br />

reported in Indonesia (Santoso 1989, Santoso et al.<br />

1989). However, it is not clear whether ectomycorrhizas<br />

were <strong>for</strong>med by the test fungi or by contaminants.<br />

In a study <strong>of</strong> site characteristics and distribution <strong>of</strong><br />

tree species in mixed dipterocarp <strong>for</strong>ests in Sarawak,<br />

Baillie and co-workers (1987) considered phosphorus<br />

the most critical nutrient while magnesium was thought<br />

to be important because <strong>of</strong> possible effects on the<br />

efficiency <strong>of</strong> the dipterocarp mycorrhizal root systems.<br />

Some species <strong>of</strong> <strong>dipterocarps</strong>, e.g. Shorea parvifolia<br />

were consistently associated with sites <strong>of</strong> high P status<br />

while others like S. quadrinervis were associated with<br />

sites <strong>of</strong> low P status. Amir and Miller (1990) found<br />

potassium to be the primary limiting nutrient in two<br />

separate <strong>for</strong>est reserves in Peninsular Malaysia. Burslem<br />

et al. (1994), however, are <strong>of</strong> the opinion that any <strong>of</strong> the<br />

macronutrients and micronutrients can become<br />

potentially limiting to plant growth when the primary<br />

limitation by P is overcome. From a study <strong>of</strong> soils under<br />

mixed dipterocarp <strong>for</strong>est in Brunei, Takahashi et al.<br />

(1994) stated that logged over <strong>for</strong>ests are suitable <strong>for</strong><br />

106<br />

enrichment planting with <strong>dipterocarps</strong> since loss <strong>of</strong> soil<br />

nutrients and degradation <strong>of</strong> nutrient status would be small<br />

because <strong>of</strong> nutrient accumulation in the deeper horizons.<br />

It is known that different tree species have differing<br />

site requirements reflecting their differing abilities to<br />

take up nutrients from intractable soil sources due to<br />

differences in root system architecture and in the<br />

particular differences in the mycorrhizal relationships<br />

between species (Miller 1991). Yasman (1995) found<br />

that light demanding Shorea leprosula seedlings could<br />

<strong>for</strong>m many more ectomycorrhizal types than shade<br />

tolerant Dipterocarpus confertus seedlings. Mineral<br />

nutrition, plant light requirements and mycorrhizal<br />

infection are very intimately related but it is only recently<br />

that the importance <strong>of</strong> this relationship has begun to<br />

receive recognition. Newton and Pigott (1991a) working<br />

with oak and birch found that application <strong>of</strong> fertilisers<br />

could reduce the number <strong>of</strong> mycorrhizal types and their<br />

relative abundances. Lee and Alexander (1994) found that<br />

full nutrient application prevented ectomycorrhizal<br />

<strong>for</strong>mation in Hopea odorata but not in H. helferi. This<br />

may indirectly affect the drought tolerance <strong>of</strong> the host<br />

plants and consequently have implications on <strong>for</strong>est<br />

management. Burslem et al. (1994) suggested that<br />

mycorrhizas play an important role in enabling<br />

Melastoma to grow on very nutrient poor soils despite<br />

being highly nutrient demanding. They suggested that <strong>for</strong><br />

mycorrhizal plants, limitation by the major cations may<br />

prove more significant than limitation by P. In a more<br />

recent study, Burslem et al. (1995) suggest that shade<br />

tolerant seedlings <strong>of</strong> lowland tropical <strong>for</strong>est which<br />

possess mycorrhizas are not limited by P supply because<br />

the mycorrhizas effectively relieve them <strong>of</strong> P limitation<br />

and/or because such plants have a low demand <strong>for</strong><br />

nutrients <strong>for</strong> growth at low irradiance.<br />

It is clear that there is an urgent need <strong>for</strong> more<br />

integrated studies on dipterocarp mineral nutrient<br />

requirements and that such studies must take into<br />

consideration the role <strong>of</strong> the dipterocarp mycorrhizal<br />

association and the effect <strong>of</strong> different light regimes.<br />

While such studies are more easily conducted in the<br />

nursery with potted plants, there is also a need to test<br />

the conclusions <strong>of</strong> such experiments in the field.<br />

<strong>Research</strong> Priorities<br />

The need <strong>for</strong> more research into the dipterocarp<br />

mycorrhizal association is already well recognised and

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