Growth of Seedlings

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).


Adams WW, Demmig-Adams B, Logan BA, Barker DH, Osmond CB (1999) Rapid changes in xanthophyll cycle-dependent energy dissipation and photosystem II efficiency in two vines, Stephania japonica and Smilax australis, growing in the understory of an open Eucalyptus forest. Plant Cell Environ 22:125-136

Agyeman VK, Swaine MD, Thompson J (1999) Responses of tropical forest tree seedlings to irradiance and the derivation of a light response index. J Ecol 87:815-827 Anderson JM, Thomson WW (1989) Dynamic molecular organization of the plant thylakoid membrane. Photosynthesis. Alan R Liss, New York, pp 161-182 Aphalo PJ, Ballaré CL, Scopel AL (1999) Plant-plant signalling, the shade-avoidance response and competition. J Exp Bot 50:1629-1634 Augustynowicz J, Gabrys H (1999) Chloroplast movements in fern leaves: correlation of movement dynamics and environmental flexibility of the species. Plant Cell Environ 22:1239-1248 Bailey S, Walters RG, Jansson S, Horton P (2001) Acclimation of Arabidopsis thaliana to the light environment: the existence of separate low light and high light responses. Planta 213:794-801 Baroli I, Do AD, Yamane T, Niyogi KK (2003) Zeaxanthin accumulation in the absence of a functional xanthophyll cycle protects Chlamydomonas reinhardtii from photooxidative stress. Plant Cell 15:992-1008

Barth C, Krause GH, Winter K (2001) Responses of photosystem I compared with photosystem II

to high-light stress in tropical shade and sun leaves. Plant Cell Environ 24:163-176 Bilger W, Bjorkman O (1994) Relationships among violaxanthin deepoxidation, thylakoid membrane conformation, and non-photochemical chlorophyll fluorescence quenching in leaves of cotton (Gossypium hirsutum L.). Planta 193:238-246 Bilger W, Schreiber U, Bock M (1995) Determination of the quantum efficiency of photosystem II and of non-photochemical quenching of chlorophyll fluorescence in the field. Oecologia 102:425-432

Bjorkman O, Demmig B (1987) Photon yield of O2 evolution and chlorophyll fluorescence characteristics at 77 K among vascular plants of diverse origins. Planta 170:489-504 Blumwald AG, Peart DR (2001) Growth strategies of a shade-tolerant tropical tree: the interactive effects of canopy gaps and simulated herbivory. J Ecol 89:608-615 Bonal D, Barigah TS, Granier A, Guehl JM (2000) Late-stage canopy tree species with extremely low S 13C and high stomatal sensitivity to seasonal soil drought in the tropical rainforest of French Guiana. Plant Cell Envoiron 23:445-459 Büch K, Stransky H, Bigus H-J, Hager A (1994) Enhancement by artificial electron acceptors of thylakoid lumen acidification and zeaxanthin formation. J Plant Physiol 144:641-648 Bungard RA, Press C, Scholes JD (2000) The influence of nitrogen on rain forest dipterocarp seedlings exposed to a large increase in irradiance. Plant Cell Environ 23:1183-1194 Castro Y, Fetcher N, Fernández DS (1995) Chronic photoinhibition in seedlings of tropical trees.

Physiol Plant 94:560-565 Cheng L (2003) Xanthophyll cycle pool size composition in relation to the nitrogen content of apple leaves. J Exp Bot 54:385-393 Chow WS, Qian L, Goodchild DJ, Anderson JM (1988) Photosynthetic acclimation of Alocasia macrorrhiza (L.) G. Don. to growth irradiance: structure, function and composition of chloro-plasts. Aust J Plant Physiol 15:107-122 Critchley C, Russell AW (1994) Photoinhibition of photosynthesis in vivo: the role of protein turnover in photosystem II. Physiol Plant 92:188-196 Demmig-Adams B (1990) Carotenoids and photoprotection: a role for the xanthophyll zeaxanthin cycle. Biochim Biophys Acta 1020:1-24 Demmig-Adams B, Adams WW (1992) Photoprotection and other responses of plants to high light stress. Annu Rev Plant Physiol Plant Mol Biol 43:599-626 Demmig-Adams B, Adams WW (1993) The xanthophyll cycle, protein turnover and the high light tolerance of sun-acclimated leaves. Plant Physiol 103:1413-1420 Demmig-Adams B, Adams WW (1994) Capacity for energy dissipation in the pigment bed in leaves with different xanthophyll cycle pools. Aust J Plant Physiol 21:575-588 Demmig-Adams B, Gilmore AM, Adams WW (1996) In vivo functions of carotenoids in higher plants. FASEB J 10:403-412 Doley D, Unwin GL, Yates DJ (1988) Spatial and temporal distribution of photosynthesis and transpiration by single leaves in a rainforest tree, Argyrodendron peralatum. Aust J Plant Physiol 15:317-326

Eschenbach C, Glauner R, Kleine M, Kappen L (1998) Photosynthesis of selected tree species in lowland dipterocarp forest of Sabah, Malaysia. Trees 12:356-365 Evans JR (1988) Acclimation by the thylakoid membranes to growth irradiance and the partitioning of nitrogen between soluble and thylakoid proteins. Aust J Plant Physiol 15:93-106 Fetene M, Lee HSJ, Lüttge U (1990) Photosynthetic acclimation in a terrestrial CAM bromeliad,

Bromelia humilis Jacq. New Phytol 114:399-406 Field CB (1988) On the role of photosynthetic responses in constraining the habitat distribution of rainforest plants. Aust J Plant Physiol 15:343-358 Flores S (1992) Growth and seasonality of seedlings and juveniles of primary species of a cloud forest in northern Venezuela. J Trop Ecol 8:299-305 Genty B, Briantais J-M, Baker NR (1989) The relationship between the quantum yield of photosynthetic electron transport and quenching of chlorophyll fluorescence. Biochin Biophy Acta 990:87-92

Gilmore AM (1997) Mechanistic aspects of xanthophyll cycle-dependent photoprotection in higher plant chloroplasts and leaves. Physiol Plant 99:197-209 Gilmore AM, Govindjee (1999) How higher plants respond to excess light: energy dissipation in photosystem II. In: Singhal GS, Renger G, Sopory SK, Irrgang K-D, Govindjee (eds) Concepts in photobiology: photosynthesis and photomorphogenesis. Narosa Publ House, New Delhi, pp 513-548

Gilmore AM, Yamasaki H (1998) 9-Aminoacridine and dibucaine exhibit competitive interactions and complicated inhibitory effects that interfere with measurements of ApH and xanthophyll cycle-dependent photosystem II energy dissipation. Photosynthesis Res 57:159-174 Gilmore AM, Hazlett TL, Debrunner PG, Govindjee (1996) Comparative time-resolved photosystem II chlorophyll a fluorescence analyses reveal distinctive differences between photoin-hibitory reaction center damage and xanthophyll cycle-dependent energy dissipation. Pho-tochem Photobiol 64:552-563 Gilmore AM, Shinkarev VP, Hazlett TL, Govindjee (1998) Quantitative analysis of the effects of intrathylakoid pH and xanthophyll cycle pigments on chlorophyll a fluorescence lifetime distributions and intensity in thylakoids. Biochemistry 73:13582-13593 Gorton H, Williams WE, Vogelmann TC (1999) Chloroplast movement in Alocasia macrorrhiza.

Physiol Plant 106:421-428 Hager A (1980) The reversible, light-induced conversions of xanthophylls in the chloroplast. In:

Czygan FC (ed) Pigments in plants. G Fischer, Stuttgart, pp 57-79 Hager A, Holocher K (1994) Localization of the xanthophyll-cycle enzyme violaxanthin de-epoxidase within the thylakoid lumen and abolition of its mobility by a (light-dependent) pH decrease. Planta 192:581-589 He J, Chow WS (2003) The rate coefficient of repair of photosystem II after photoinactivation.

Physiol Plant 118:297-304 He J, Chee CW, Goh CJ (1996) 'Photoinhibition' of Heliconia under tropical conditions: the importance of leaf orientation for light interception and leaf temperature. Plant Cell Envoiron 19:1238-1248

Holt NE, Zigmantas D, Valkunas L, Li X-P, Niyogi KK, Fleming GR (2005) Carotenoid cation formation and the regulation of photosynthetic light harvesting. Science 307:433-436 Horton P, Ruban A (2005) Molecular design of the photosystem II light-harvesting antenna: photosynthesis and photoprotection. J Exp Bot 56:365-373 Horton P, Ruban AV, Walters RG (1994) Regulation of light harvesting in green plants. Indication by nonphotochemical quenching of chlorophyll fluorescence. Plant Physiol 106:415-420 Huber O (1978) Light compensation point of vascular plants of a tropical cloud forest and an ecological interpretation. Photosynthetica 12:382-390 Huc R, Ferhi A, Guehl JM (1994) Pioneer and late stage tropical rainforest tree species (French Guiana) growing under common conditions differ in leaf gas exchange regulation, carbon isotope discrimination and leaf water potential. Oecologia 99:297-305

Humbeck K, Senger H (1984) The blue light factor in sun and shade plant adaptation. In: Senger H (ed) Blue light effects in biological systems. Springer, Berlin Heidelberg New York, pp 344-351

King DA, Davies SJ, Tan S, Noor NSMD (2006) The role of wood density and stem support costs in the growth and mortality of tropical trees. J Ecol 94:670-680 Kirschbaum MUF, Pearcy RW (1988a) Concurrent measurements of oxygen- and carbon-dioxide exchange during light flecks in Alocasia macrorrhiza (L.) G. Don. Planta 174:527-533 Kirschbaum MUF, Pearcy RW (1988b) Gas exchange analysis of the fast phase of photosynthetic induction in Alocasia macrorrhiza. Plant Physiol 87:818-821 Kirschbaum MUF, Küppers M, Schneider H, Giersch C, Noe S (1997) Modelling photosynthesis in fluctuating light with inclusion of stomatal conductance, biochemical activation and pools of key photosynthetic intermediates. Planta 204:16-26 Kitajima K (1994) Relative importance of photosynthetic traits and allocation patterns as correlates of seedling shade tolerance of 13 tropical trees. Oecologia 98:419-428 Kitajima K, Hogan KP (2003) Increases of chlorophyll a/b ratios during acclimation of tropical woody seedlings to nitrogen limitation and high light. Plant Cell Environ 26:857-865 Königer M, Harris GC, Pearcy RW (1998) Interaction between flux density and elevated temperatures on photoinhibition in Alocasia macrorrhiza. Planta 205:214-222 Koten O van, Snel JFH (1990) The use of chlorophyll fluorescence nomenclature in plant stress physiology. Photosynthesis Res 25:147-150 Krause GH, Virgo A, Winter K (1995) High susceptibility to photoinhibition of young leaves of tropical forest trees. Planta 197:583-591 Krause GH, Schmude C, Garden H, Koroleva O, Winter K (1999) Effects of solar ultraviolet radiation on the potential efficiency of photosystem II in leaves of tropical plants. Plant Physiol 121:1349-1358

Krause GH, Koroleva OY, Dalling JW, Winter K (2001) Acclimation of tropical tree seedlings to excessive light in simulated tree-fall gaps. Plant Cell Environ 24:1345-1352 Krause GH, Galle A, Gademann R, Winter K (2003a) Capacity of protection against ultraviolet radiation in sun and shade leaves of tropical forest plants. Funct Plant Biol 30:533-542 Krause GH, Grube E, Virgo A, Winter K (2003b) Sudden exposure to solar UV-B radiation reduces net CO2 uptake and photosystem I efficiency in shade-acclimated tropical tree seedlings. Plant Physiol 131:745-758

Krause GH, Grube E, Koroleva OY, Barth C, Winter K (2004) Do mature shade leaves of tropical tree seedlings acclimate to high sunlight and UV radiation? Funct Plant Biol 31:743-756 Krause GH, Gallé A, Virgo A, Garcia M, Bucic P, Jahns P, Winter K (2006) High-light stress does not impair biomass accumulation of sun-acclimated tropical tree seedlings (Calophyllum longifolium Willd. and Tectona grandis L.f.). Plant Biol 8:31-41 Krauss P, Markstätter C, Riederer M (1997) Attenuation of UV radiation by plant cuticles from woody species. Plant Cell Environ 20:1079-1085 Krieger-Liszkay A, Trebst A (2006) Tocopherol is the scavenger of singlet oxygen produced by the triplet state of chlorophyll in the PSII reaction centre. J Exp Bot 57:1677-1684 Kursar TA, Coley PD (1993) Photosynthetic induction times in shade-tolerant species with long and short-lived leaves. Oecologia 93:165-170 Kyereh B, Swaine MD, Thompson J (1999) Effect of light on the germination of forest trees in

Ghana. J Ecol 87:772-783 Leakey ADB, Press MC, Scholes JD (2003) High-temperature inhibition of photosynthesis is greater under sunflecks than uniform irradiance in a tropical rain forest tree seedling. Plant Cell Environ 26:1681-1690 Leakey ADB, Scholes JD, Press MC (2005) Physiological and ecological significance of sunflecks for dipterocarp seedlings. J Exp Bot 56:469-482 Lichtenthaler HK, Buschmann C, Döll M, Fietz H-J, Bach T, Kozel U, Meier D, Rahmsdorf U (1981) Photosynthetic activity, chloroplast ultrastructure and leaf characteristics of high-light and low-light plants and of sun and shade leaves. Photosynth Res 2:115-141

Logan BA, Barker DH, Demmig-Adams B, Adams WW (1996) Acclimation of leaf carotenoid composition and ascorbate levels to gradients in the light environment within an Australian rainforest. Plant Cell Environ 19:1083-1090 Logan BA, Barker DH, Adams WW, Demmig-Adams B (1997) The response of xanthophyll cycle-dependent energy dissipation in Alocasia brisbanensis to sunflecks in a subtropical rainforest. Aust J Plant Physiol 24:27-53 Lüttge U (1985) Epiphyten:Evolution und Ökophysiologie. Naturwissenschaften 72:557-566 Lüttge U, Ball E, Kluge M, Ong BL (1986) Photosynthetic light requirements of various tropical vascular epiphytes. Physiol Vég 24:315-331 Lüttge U, Berg A, Fetene M, Nauke P, Peter D, Beck E (2003) Comparative characterization of photosynthetic performance and water relations of native trees and exotic plantation trees in an Ethiopian forest. Trees 17:40-50 Lüttge U, Kluge M, Bauer G (2005) Botanik, 5th edn. VCH, Weinheim

Matsubara S, Gilmore AM, Osmond CB (2001) Diurnal and acclimatory responses of violaxanthin and lutein epoxide in the Australian mistletoe Amyema miquelii. Aus J Plant Physiol 28:793800

Matsubara S, Morosinotto T, Bassi R, Christian A-L, Fischer-Schliebs E, Lüttge U, Orthen B, Franco AC, Scarano FR, Förster B, Pogson BJ, Osmond CB (2003) Occurrence of the lutein-epoxide cycle in mistletoes of the Loranthaceae and Viscaceae. Planta 217:868-879 Matsubara S, Naumann M, Martin R, Nichol C, Rascher U, Morosinotto T, Bassi R, Osmond B (2005) Slowly reversible de-epoxidation of lutein-epoxide in deep shade of a tropical tree legume may 'lock-in' lutein-based photoprotection during acclimation to strong light. J Exp Bot 56:461-468

Maxwell K, Johnson GN (2000) Chlorophyll fluorescence - a practical guide. J Exp Bot 51:659668

Maxwell C, Griffiths H, Young AJ (1994) Photosynthetic acclimation to light regime and water stress by the C3-CAM epiphyte Guzmania monostachia: gas exchange characteristics, photochemical efficiency and the xanthophyll cycle. Funct Ecol 8:746-754 Maxwell C, Griffiths H, Borland AM, Young AJ, Broadmeadow MSJ, Fordham MC (1995) Short-term photosynthetic responses of the C3-CAM epiphyte Guzmania monostachia var. monos-tachia to tropical seasonal transitions under field conditions. Aust J Plant Physiol 22:771-781 Medina E (1998) Seedling establishment and endurance in tropical forests: ecophysiology of stress during early stages of growth. In: Scarano FR, Franco AC (eds) Ecophysiological strategies of xerophytic and amphibious plants in the neotropics. Oecologia Brasiliensis, vol IV. PPGE-UFRJ, Rio de Janeiro, pp 23-43 Myers JA, Kitajima K (2007) Carbohydrate storage enhances seedling shade and stress tolerance in a neotropical forest. J Ecol 95:383-395 Newell EA, McDonald EP, Strain BR, Denslow JS (1993) Photosynthetic responses of Miconia species to canopy openings in a lowland tropical rainforest. Oecologia 94:49-56 Nielsen SL, Enriquez S, Duarte CM (1997) Control of PAR-saturated CO2 exchange rate in some

C3 and CAM plants. Biol Plant 40:91-101 Noguchi K, Nakajima N, Terashima I (2001a) Acclimation of leaf respiratory properties in Alocasia odora following reciprocal transfers of plants between high- and low-light environments. Plant Cell Environ 24:831-839 Noguchi K, Go C-S, Terashima I, Ueda S, Yoshinari T (2001b) Activities of the cyanide-resistant respiratory pathway in leaves of sun and shade species. Aust J Plant Physiol 28:27-35 Osmond CB, Grace SC (1995) Perspectives on photoinhibition and photorespiration in the field: quintessential inefficiencies of the light and dark reactions of photosynthesis? J Exp Bot 46:1351-1362

Pearcy RW (1983) The light environment and growth of C3 and C4 tree species in the understory of a Hawiian forest. Oecologia 58:19-25 Pearcy RW, Osteryoung K, Calkin HW (1985) Photosynthetic responses to dynamic light environments by Hawaiian trees. Plant Physiol 79:896-902

Pfündel E, Bilger W (1994) Regulation and possible function of the violaxanthin cycle. Photosynth Res 42:89-109

Poorter L (1998) Seedling growth of Bolivian rain forest species in relation to light and water availability. PhD Thesis, Utrecht (ISBN 90-393-1759-3) Poorter L, Oberbauer SF (1993) Photosynthetic induction responses of two rainforest tree species in relation to light environment. Oecologia 96:193-199 Poorter L, Bongers F, Sterck FJ, Wöll H (2005) Beyond the regeneration phase: differentiation of height-light trajectories among tropical tree species. J Ecol 93:256-267 Ramalho JC, Compos PS, Quartin VL, Silva MJ, Nunes MA (1999) High irradiance impairments on photosynthetic electron transport, ribulose-1,5-bisphosphate carboxylase/oxygenase and N assimilation as a function of N availability in Coffea arabica L. plants. J Plant Physiol 154:319-326

Reich PB, Walters MB, Ellsworth DS, Uhl C (1994) Photosynthesis-nitrogen relations in Amazonian tree species. I. Patterns among species and communities. Oecologia 97:62-72 Riddoch I, Lehto T, Grace J (1991) Photosynthesis of tropical tree seedlings in relation to light and nutrient supply. New Phytol 119:137-147 Sack L, Tyree MT, Holbrook NM (2005) Leaf hydraulic architecture correlates with regeneration irradiance in tropical rainforest trees. New Phytol 167:403-413 Sassenrath-Cole GF, Pearcy RW (1992) The role of ribulose-1,5-bisphosphate regeneration in the induction requirement of photosynthetic CO2 exchange under transient light conditions. Plant Physiol 99:227-234

Schäfer C, Schmid V (1993) Pflanzen im Lichtstreß. Biol Unserer Zeit 23:55-62 Schiefthaler U, Russell AW, Bolhar-Nordenkamf HR, Critchley C (1999) Photoregulation and photodamage in Schefflera arboricola leaves adapted to different light environments. Aust J Plant Physiol 26:485-494

Schindler C, Lichtenthaler HK (1996) Photosynthetic CO2-assimilation, chlorophyll fluorescence and zeaxanthin accumulation in field grown maple trees in the course of a sunny and a cloudy day. J Plant Physiol 148:399-412 Scholes JD, Press MC, Zipperlen SW (1997) Differences in Light energy utilization and dissipation between dipterocarp forest tree seedlings. Oecologia 109:41-48 Schreiber U, Bilger W (1987) Rapid assessment of stress effects on plant leaves by chlorophyll fluorescence measurements. In: Tenhunen JD, Catarino FM, Lange OL, Oechel WC (eds) Plant responses to stress. Functional analysis in Mediterranean ecosystems. NATO-ASI-Series G, Ecological sciences, vol 15. Springer, Berlin Heidelberg New York, pp 27-53 Schreiber U, Bilger W (1993) Progress in chlorophyll fluorescence research: major developments during the past years in retrospect. Prog Bot 54:151-173 Sims DA, Pearcy RW (1994) Scaling sun and shade photosynthetic acclimation of Alocasia macr-orrhiza to whole-plant performance. - I. Carbon balance and allocation at different daily photon flux densities. Plant Cell Environ 17:881-887 Sims DA, Gebauer RLE, Pearcy RW (1994) Scaling sun and shade photosynthetic acclimation of Alocasia macrorrhiza to whole-plant performance. - II. Simulation of carbon balance and growth at different photon flux densities. Plant Cell Environ 17:889-900 Smith H, Samson G, Fork DC (1993) Photosynthetic acclimation to shade:probing the role of phytochromes using photomorphogenic mutants of tomato. Plant Cell Environ 16:929-937 Stegemann J, Timm H-C, Küppers M (1999) Stimulation of photosynthetic plasticity in response to high fluctuating light: an empirical model integrating dynamic photosynthetic induction and capacity. Trees 14:145-160 Stephens GR, Waggoner PE (1970) Carbon dioxide exchange of a tropical rainforest. Part I. BioScience 20:1050-1053

Strauss-Debenedetti S, Bazzaz FA (1991) Plasticity and acclimation to light in tropical Moraceae of different successional positions. Oecologia 87:377-387 Thiele A, Winter K, Krause GH (1997) Low inactivation of D1 protein of photosystem II in young canopy leaves of Anacardium excelsum under high-light stress. J Plant Physiol 151:286-292

Thiele A, Krause GH, Winter K (1998) In situ study of photoinhibition of photosynthesis and xanthophyll cycle activity in plants growing in natural gaps of the tropical forest. Aust J Plant Physiol 25:189-195

Timm H-C, Stegemann J, Küppers M (2002) Photosynthetic induction strongly affects the light compensation point of net photosynthesis and coincidentally the apparent quantum yield. Trees 16:47-62

Timm H-C, Küppers M, Stegemann J (2004) Non-destructive analysis of architectural expansion and assimilate allocation in different tropical tree saplings: consequences of using steady-state and dynamic photosynthesis models. Ecotropica 10:101-121 Tinoco-Ojanguren C, Pearcy RW (1992) Dynamic stomatal behaviour and its role in carbon gain during lightflecks of a gap phase and an understory Piper species acclimated to high and low light. Oecologia 92:222-228 Tinoco-Ojanguren C, Pearcy RW (1993a) Stomatal dynamics and its importance to carbon gain in two rainforest Piper species. I. VPD effects on the transient stomatal response to light flecks. Oecologia 94:388-394

Tinoco-Ojanguren C, Pearcy RW (1993b) Stomatal dynamics and its importance to carbon gain in two rainforest Piper species. II. Stomatal versus biochemical limitations during photosynthetic induction. Oecologia 94:395-402 Toth VR, Meszäros I, Veres S, Nagy J (2002) Effects of the available nitrogen on the photosynthetic activity and xanthophyll cycle pool of maize in the field. J Plant Physiol 159:627-634 Turnball MH (1991) The effect of light quantity and quality during development on the photosyn-

thetic characteristics of six Australian rainforest tree species. Oecologia 87:110-117 Tyystjärvi E, Aro E-M (1996) The rate constant of photoinhibition measured in lincomycin-treated leaves is directly proportional to light intensity. Proc Nat Acad Sci USA 93:2213-2218 Valladares F, Allen MT, Pearcy RW (1997) Photosynthetic responses to dynamic light under field conditions in six tropical rainforest shrubs occurring along a light gradient. Oecologia 111:505-514

Väzquez-Yanes C, Orozco-Segovia A (1993) Patterns of seed longevity and germination in the tropical rainforest. Annu Rev Ecol Syst 24:69-87 Vogelmann TC, Bornman JF, Yates DJ (1996) Focussing of light by leaf epidermal cells. Physiol Plant 98:43-56

Warren CR, Adams MA, Chen ZL (2000) Is photosynthesis related to concentration of nitrogen and Rubisco in leaves of Australian native plants? Aust J Plant Physiol 27:407-416 Watling JR, Robinson SA, Woodrow IE, Osmond CB (1997) Responses of rainforest understory plants to excess light during sunflecks. Aust J Plant Physiol 24:17-25 Yamashita N, Koike N, Ishida A (2002) Leaf ontogenetic dependence of light acclimation in invasive and native subtropical trees of different successional status. Plant Cell Environ 25:13411356

Zipperlen SW, Press MC (1996) Photosynthesis in relation to growth and seedling ecology of two dipterocarp rain forest tree species. J Ecol 84:863-876 Zipperlen SW, Press MC (1997) Photosynthetic induction and stomatal oscillations in relation to the light environment of two dipterocarp rain forest tree species. J Ecol 85:491-503

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