Seedling growth depends on availability of water (Sect. 5.1) and nutrients (Sect. 3.4.4) but most drastically on light (Medina 1998; Poorter 1998). After germination and establishment of seedlings often forming nurseries of slow growing plants (Sect. 4.3.1), further growth will depend very much on photosynthesis, the light powered fixation of CO2 and reduction to organic material, as well as allocation of photosynthetic products (Zipperlen and Press 1996; Scholes et al. 1997). This, of course, applies to both pioneer and climax species. Both may differ though, in responses to light intensity, which are representative of sun and shade plant characteristics (see Sect. 4.1.1).
However, in some cases the differences in photosynthetic capacity related to light intensity between seedlings of pioneer and climax species, or between mature early and late successional species, has been found to be surprisingly small (Riddoch
Fig. 4.15 Seedling mortality rate during the first year in the shade of 13 tropical tree species related to relative growth rate (RGR) in shade and sun (A,B), leaf area ratio (LAR) (C), root : shoot ratio (D) and wood density (E). (After Kitajima 1994)
Fig. 4.16 Leaf-anatomy of seedlings of a pioneer or early succession tree, Nau-clea diderrichii (A,B), and a late succession tree Entan-drophragma angolense (C,D), grown at high light (A,C) and at low light (B,D). The white bars denote 50 ^ m. (Riddoch etal. 1991)
et al. 1991). Huber (1978) examined photosynthetic characteristics, e.g. the light compensation point (Sect. 4.1.1), of 54 vascular plant species in Rancho Grande (Venezuela). He found that by this criterion the majority of the species growing in the lower forest strata did not belong to extreme shade-adapted plant types, but possessed a wide capacity for response to the highly variable irradiance in this montane cloud forest. It may be noted generally that a schematic distinction of pioneer sun plants/shade plants in this context is too simple. Changes can occur during development (Turnball 1991; Agyeman et al. 1999; Kyereh et al. 1999; Poorter et al. 2005). Mature shade leaves of seedlings can substantially acclimate to full sunlight employing mechanisms of energy dissipation, UV absorbing substances etc. (Krause et al. 2004). Availability of water and nutrients, especially nitrogen (Sect. 4.1.2) play a role in this (Castro et al. 1995; Bungard et al. 2000).
Complex regulation is involved. For a fluctuating tropical environment with frequent disturbance by typhoons and canopy opening a trade off between acclimation ability and plasticity has been considered (Yamashita et al. 2002). Strauss-Debenedetti and Bazzaz (1991) have suggested that plasticity and acclimation should be distinguished as follows:
• late successional species often cannot acclimate to high light intensities when transferred from low-light to high light (low acclimation) but may grow well if kept continuously under low and high light respectively (high plasticity),
• pioneer species may grow at low and high light and show a considerable stimulation after transfer from low light to high light (high plasticity and high acclimation).
The expression of low-light and high-light forms of a species may also be determined by the phytochrome system (Smith et al. 1993), but in particular blue-light photoreceptors are also involved in this regulation (Lichtenthaler et al. 1981; Humbeck and Senger 1984; see Luttge et al. 1986). Leaf-anatomical features often show pronounced differences; sun leaves are thicker than shade leaves and have additional layers of palisade parenchyma. In a comparison of young seedlings of the tropical trees Nauclea diderrichii (De Wilde.) Merrill, a pioneer species, and En-tandrophragma angolense (Welw.) C.DC., a climax species, both from West Africa, differences in acclimation and photosynthetic capacity at high light intensity were only small. However, there were marked morphogenetic effects on leaf anatomy in plants grown in the sun and in the shade respectively, in the pioneer species N. diderrichii but not so much in the climax species E. angolense (Fig. 4.16).
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