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A review of dipterocarps - Center for International Forestry Research

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Root Symbiosis and Nutrition<br />

However, no data on the nature or level <strong>of</strong> mycorrhizal<br />

infection were presented. The report by Louis and Scott<br />

(1987) <strong>of</strong> mycorrhizal synthesis in root organ cultures<br />

<strong>of</strong> Shorea roxburghii can be discounted as their<br />

illustrations and descriptions do not show<br />

ectomycorrhizas but hyphal invasion into root cells.<br />

Moreover the fungus they used, Rhodophyllus sp. was<br />

from a genus not normally considered to be<br />

ectomycorrhizal.<br />

The effects <strong>of</strong> various environmental factors on<br />

dipterocarp ectomycorrhizas and their subsequent<br />

effects on plant growth have been the subject <strong>of</strong> some<br />

recent studies. Smits (1994) suggested that the obligate<br />

nature and temperature sensitivity <strong>of</strong> the dipterocarp<br />

ectomycorrhizal relationship are the determining factors<br />

<strong>for</strong> good dipterocarp seedling per<strong>for</strong>mance. Yasman<br />

(1995) stated that light intensity influenced<br />

ectomycorrhizal <strong>for</strong>mation in dipterocarp seedlings but<br />

that the effects varied between different host species.<br />

The physiology <strong>of</strong> how light regulated ectomycorrhizal<br />

<strong>for</strong>mation was, however, not examined. According to<br />

Yasman (1995), neither light nor soil conditions<br />

represented the main factors <strong>for</strong> successful dipterocarp<br />

regeneration under a closed canopy; dipterocarp seedling<br />

survival was mainly related to the presence <strong>of</strong><br />

mycorrhizal inoculum and the support <strong>of</strong> the seedlings<br />

by their ectomycorrhizal connections to roots from<br />

mother trees that had well illuminated emergent crowns.<br />

However, this may be an oversimplification as different<br />

species <strong>of</strong> <strong>dipterocarps</strong> have different light requirements<br />

(Mori 1980, Sasaki and Mori 1981). Lee et al. (in press)<br />

found high levels <strong>of</strong> ectomycorrhizal infection (60%)<br />

on seedlings <strong>of</strong> Hopea nervosa and Shorea leprosula<br />

under heavy shade in undisturbed <strong>for</strong>est supporting<br />

Yasman’s (1995) hypothesis, but also found that S.<br />

leprosula which is a light demanding species had poor<br />

survival compared to H. nervosa which is a shade<br />

tolerant species.<br />

Mineral Nutrition<br />

It must be emphasised that very few studies have been<br />

conducted on the very important aspect <strong>of</strong> mineral<br />

nutrient requirements <strong>of</strong> <strong>dipterocarps</strong>. Fertiliser trials<br />

have been conducted <strong>for</strong> several dipterocarp species but<br />

the in<strong>for</strong>mation presently available is far from<br />

conclusive. Although a preliminary guide <strong>for</strong> the<br />

diagnosis <strong>of</strong> nutrient deficiency <strong>of</strong> tropical <strong>for</strong>est trees<br />

105<br />

has been developed (Fassbender 1988), its applicability<br />

and suitability <strong>for</strong> <strong>dipterocarps</strong> has to be tested more<br />

extensively.<br />

Sundralingam (1983) found that NP fertilisers<br />

generally improved growth <strong>of</strong> potted Dryobalanops<br />

aromatica and D. oblongifolia seedlings. In another<br />

experiment, Sundralingam and her co-workers (1985)<br />

found that nitrogen rather than phosphorus was the most<br />

important fertiliser required <strong>for</strong> improved growth <strong>of</strong><br />

potted Shorea ovalis seedlings. Madius (1983) found<br />

that potted Shorea bracteolata and S. parvifolia<br />

seedlings had improved growth and increased nutrient<br />

uptake at higher fertiliser levels, particularly when<br />

moisture supply was abundant. Turner et al. (1993),<br />

however, found that potted Shorea macroptera seedlings<br />

did not respond to fertiliser application. However, they<br />

found that the extent <strong>of</strong> mycorrhizal infection on S.<br />

macroptera seedlings was correlated with seedling dry<br />

mass in the unfertilised control. Similarly, Burslem et<br />

al. (1995) working with potted Dipterocarpus kunstleri<br />

seedlings, found no positive growth response to nutrient<br />

additions although addition <strong>of</strong> P increased the<br />

concentrations <strong>of</strong> K and Ca in the leaves. Burslem and<br />

his co-workers (1995) caution that results <strong>of</strong> pot<br />

bioassay experiments may be dependent on factors such<br />

as pot size, irradiance and soil moisture conditions and<br />

that conclusions drawn from such experiments need to<br />

be tested by field fertilisation experiments.<br />

Turner et al. (1993) also reported that naturally<br />

regenerated seedlings <strong>of</strong> Shorea curtisii and Hopea<br />

beccariana did not show any significant response to<br />

fertiliser application in the field. They suggested that<br />

addition <strong>of</strong> nutrients to promote higher growth rates in<br />

regenerating seedlings in dipterocarp <strong>for</strong>ests is unlikely<br />

to be a silvicultural practice although ensuring adequate<br />

infection could be beneficial. Aminah and Lokmal (1995)<br />

reported that outplanted rooted, stem cuttings <strong>of</strong> H.<br />

odorata showed a significant increase in stem diameter<br />

and height only nine to 24 months after application <strong>of</strong><br />

granular compound fertiliser. No growth response was<br />

recorded when plants were measured earlier. Nussbaum<br />

et al. (1994) found that nutrient availability was the major<br />

factor limiting the establishment <strong>of</strong> Dryobalanops<br />

lanceolata and S. leprosula seedlings on degraded soils<br />

in Sabah rather than low soil moisture or high soil<br />

temperature. Preliminary results <strong>of</strong> the effects <strong>of</strong> NPK<br />

fertilisers on the growth <strong>of</strong> D. lanceolata and S.<br />

leprosula on a large-scale enrichment planting project

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