The structuring role of free-floating versus submerged plants in a subtropical shallow lake

In shallow temperate lakes many ecological processes depend on submerged macrophytes. In subtropical and tropical lakes, free-floating macrophytes may be equally or more important. We tested the hypothesis that different macrophyte growth forms would be linked with different bottom-up and top-down mechanisms in out-competing phytoplankton. We compared experimentally the effects of submerged and free-floating plants on water chemistry, phytoplankton biomass, zooplankton and fish community structure in a shallow hypertrophic lake (Lake Rodó, 34°55S 56°10W, Uruguay). Except for the retention of suspended solids, we found no other significant bottom-up process connected with either Eichhornia crassipes or Potamogeton pectinatus. Free-floating plants had a lower abundance of medium-sized zooplankton than any other microhabitat and submerged plants were apparently preferred by microcrustaceans. Fish showed a differential habitat use according to species, size-class and feeding habits. Dominant omnivore-planktivores, particularly the smallest size classes, preferred submerged plants. In contrast, omnivore-piscivores were significantly associated with free-floating plants. The density of omnivorous-planktivorous fish, by size class, significantly explained the distribution of medium-sized zooplankton, the high number of size 0 fish being the main factor. The abiotic environment and the structure of the zooplankton community explained little of the fish distribution pattern. Our results suggest that bottom-up effects of free-floating plants are weak when cover is low or intermediate. Top-down effects are complex, as effects on zooplankton and fish communities seem contradictory. The low piscivores:planktivores ratio in all microhabitats suggests, however, that cascading effects on phytoplankton through free-floating plant impacts on piscivorous fish are unlikely to be strong.     

Application of randomization methods to study the association of Trichocereus pasacana (Cactaceae) with potential nurse plants

Spatial distribution of plants and seeds of Trichocereuspasacana was studied in relation to bush species abundance, and ofcacti in relation to the available number of seeds under bush species, in LosCardones National Park, Argentina. Randomization tests, carried out to find thesignificance level of Chi squared test statistics used to test fornon-random association hypothesis, proved that T.pasacana plants, seeds, and seedling establishment is related to thedifferent bush species that co-occur with it. Positive interactions werefound mostly with L. divaricata, but also withP.ferox and A. spartioides, suggesting that theyact as nurse plants. Negative interactions were found with B.boliviensis, A. spartioides, S.crassiramea, and Verbena sp, in spite of thehigh seed storage beneath their canopies. Cacti are more frequent than expectedin relation to seed relative abundance under L. divaricataand P. ferox and less frequent under P.rougesii, B. boilviensis,Verbena sp, G. glutinosa,S.crassiramea, and A. spartioides, due todifferential seed or seedling mortality when emergence occurs.     

The shrub Rhodomyrtus tomentosa acts as a nurse plant for seedlings differing in shade tolerance in degraded land of South China

Questions: What are the nurse effects of Rhodomyrtus tomentosa in degraded land of South China? Are canopy or soil factors responsible for the main nurse effect? Do facilitative effects increase with the shade tolerance of the target species? Location: Degraded shrubland in South China. Methods: Seedlings of three native climax woody species (Schima superba, Michelia macclurei, Castanopsis fissa) that differ in shade tolerance were subjected to four treatments by transplantation: (1) under the canopy of R. tomentosa shrubs; (2) in open interspaces without vegetation cover (control); (3) under the canopy of R. tomentosa from which canopies had been removed; and (4) in open interspaces without vegetation but covered by branches and leaves of R. tomentosa. Results: At low soil nutrient levels, increased canopy shade, soil porosity and soil moisture under the canopy of R. tomentosa increased seedling survival of the climax tree species S. superba, C. fissa and M. macclurei, and shoot height of S. superba. The nurse effect (a form of facilitation) of R. tomentosa depended more on canopy shade than on soil amelioration. The magnitude of facilitation or nurse effect was positively correlated with shade tolerance of the target species. Conclusions: Use of nurse plants in restoration differs from traditional reforestation (clearing and/or burning to reduce interspecific competition between target tree species and non‐target species) because it focuses on positive interactions between nurse plants and target plants that increase establishment of target species and reduce time required for restoration. Because nurse effects of R. tomentosa shrubs tended to be larger on target species with greater shade tolerance, shade‐tolerant plants are suggested as target species to accelerate restoration.     

Biomass allocation in herbaceous plants under grazing impact in the high semi-arid Andes

Among the plant traits that affect performance, vitality and herbivore resistance in rangeland vegetation, biomass partitioning ranks top, commonly outweighing processes at single leaves (e.g., photosynthesis, respiration). We explored the allometry of a broad sample of herbaceous species from two high elevation sites in the Andes in order to explain biomass partitioning under harsh climatic conditions and risk of biomass losses under strong camelid grazing pressure. We combined data from NW-Argentina and W-Bolivia from elevations between 4200 and 4250 m in a landscape dominated by tall Festuca orthophylla tussocks and a drought driven seasonality (rainfall only between November and March). Across 10–20 taxa per region we found less investment in leaves with a mean leaf mass fraction of only 11% and instead a massive storage in below-ground compartments (rhizomes, tap roots), particularly at the colder Argentinean site. Though grazing pressure was much greater in Bolivia, the foliage mass fraction was there larger than in Argentina. The inter-tussock space in these open, dry plains (‘pajonal’) was dominated by rosette forming species with a below-ground shoot apex and massive tap roots (70% of all species), rendering these species less sensitive to grazing and trampling. The storage organs of these species represented more than 50% of total biomass. Llamas, which represented the main vertebrate herbivore in these open plains at the Bolivian site, preferred non tap-root herbs and species with low leaf nitrogen concentration. Palatable forbs for llamas (22% of all species at the Bolivian site) only survived when nested (facilitated) in the rigid, tall Festuca orthophylla tussocks or thorny shrubs. In conclusion, these extremely high elevation rangeland herbs invest preferentially in structures for persistence (K-strategy) rather than maximizing carbon gain.     

The role of cotyledons in the establishment of Suaeda physophora seedlings

The role of cotyledons in seedling establishment of the euhalophyte Suaeda physophora under non-saline and saline conditions (addition of 1 mM or 400 mM NaCl) was investigated. Survival and fresh and dry weights were greater for seedlings grown in the light (12-h light/12-h dark) than in the dark (24-h dark). The shading of cotyledons tended to decrease shoot height, shoot organic dry weight, number of leaves, and survival of seedlings regardless of NaCl treatment, but the effect of cotyledon shading was greater with 400 mM NaCl. Concentrations of Na⁺ were higher in cotyledons than in leaves, regardless of NaCl treatment. The K⁺/Na⁺ ratio was lower in cotyledons than in leaves for seedlings treated with 1 mM NaCl but not for seedlings treated with 400 mM NaCl. Addition of 400 mM NaCl decreased oxygen production in cotyledons but especially in leaves. These results are consistent with the hypothesis that, by generating oxygen via photosynthesis and by compartmentalizing Na⁺, cotyledons are crucial for the establishment of S. physophora seedlings in saline environments.     

Presence of cushion plants increases community diversity in the high equatorial Andes

Cushion plants are a common growth form in the equatorial páramo vegetation and their surfaces are often colonized by other plants. This paper analyzes the effect of the cushion plants on the community diversity at 4650 m on the eastern slope of the Iliniza volcano in Ecuador. Ninety sample plots of 1 m² size were located in the study area and were divided into 25 subplots in which presence and abundance of plant species was recorded. The community diversity was expressed as species richness, Simpson's diversity index, and evenness. Correlation between the cushion species and the composition of the colonists was measured with the CCA ordination analysis, correlation between the cushion cover and community diversity was measured by means of correlation analysis. Randomized species–area curves were used to compare richness of plant communities with and without the cushions. A total of 32 species were found including five cushion plants. Most species preferred to grow on the cushion surface whereas only a few species were able to colonize open ground. Species richness and Simpson's index were significantly correlated to the cushion area but no correlation was found for evenness. The cushions were usually composed of more than one species which hampered the examination of the cushion–colonist specific relationships. Nevertheless, cushions of Azorella and Arenaria seemed to provide more favorable habitat for colonization than the other cushion species. Comparison with an earlier study made on Iliniza indicates that the presence of the cushions significantly increases the richness of the plant community.     

Rapid root extension during water pulses enhances establishment of shrub seedlings in the Atacama Desert

Questions: (1) What is the water threshold for shrub seedling establishment in arid scrubland? (2) How do seedling root growth morphological traits affect the water threshold required for seedling establishment? Location: Arid scrubland, Atacama Desert, north‐central Chile. Methods: We conducted a field experiment with nine native shrub species under a gradient of simulated rainfall to test if species with deep root architecture have higher seedling survival rates and establish more successfully during water pulses. Results: Seedling survival rate was very low; only 2% of the 12 150 planted seedlings survived the first summer drought. Seedling survival required at least 206 mm of water, which is twice the average rainfall and roughly equivalent to the precipitation during an ENSO (El Niño Southern Oscillation) event in this region. Seedling survival at the onset of the summer drought was best explained by leaf habit, root length and initial seedling size. However, only Senna cumingii seedlings survived through the first year. S. cumingii seedlings had distinctively longer roots than the other shrub species, enabling them to reach moister soil layers. Conclusions: Water conditions resembling rainy years enhance shrub seedling establishment in the Atacama Desert, but the effects of higher water availability are strongly dependent on the shrub species. Rapid and deep rooting appears to be a very important functional trait for successful first‐year survival in this arid system where water availability is highly episodic.     

Facilitation of seedling establishment by an endemic shrub in tropical coastal sand dunes

Facilitation is predicted to occur on coastal sand dunes as these ecosystems have the harsh physical conditions common during primary succession. In a tropical dune system on the Gulf of Mexico the spatial patterns of plant distribution were analyzed and the hypothesis that facilitation is the responsible mechanism was tested by monitoring changes in the micro-environment and performing a seedling planting experiment under natural conditions. Densities of seedlings and adults of late colonizer grasses were two to six times greater beneath the early colonizer shrub, Chamaecrista chamaecristoides, than in the exposed areas. Temperatures on the sand surface, wind speed and sand accretion were significantly reduced by the shade of Chamaecrista. Only phosphate contents during the dry season were significantly higher in sand in the shade than in exposed sand. In the field experiment, successful establishment of seedlings of two late colonizer grasses (Trachypogon plumosus and Schizachyrium scoparium) was low but was concentrated exclusively beneath the shrubs. The introduced plants were reproductive one year after onset of the experiment and remained vigorous after three years. The observed spatial aggregation among the target species may be a result of improved conditions in the shade of the shrub, which facilitated the survival and establishment of late colonizers. On top of the environmental amelioration, protection from recurrent disturbances such as substrate mobility is an additional beneficial effect of the shrub.     

Spatial structure of litter-dwelling ant distribution in a subtropical dry forest

Understanding the spatial patterns of species distribution is essential to characterize the structure of communities, to optimize species inventories and to evaluate the impact of biotic and abiotic variables. Here we describe the spatial structure of the distribution of leaf litter ant species, and of biotic factors that could explain it, in a subtropical semi-deciduous forest of the Argentinian Chaco, characterized by a dense understorey of shrubs and terrestrial bromeliads. Environmental variables (leaf litter quantity and ground bromeliad density) were measured and ants were collected in 1 m² quadrats distributed along two 200 m transects at intervals of 1.25 m. Overall 87 species were collected. Sixteen positive associations and a single negative association were observed between the 11 most frequent species taken pair-wise. Our results suggest that the spatial distribution of leaf litter ants was determined at two different scales. At a small scale (period below 10 m) a periodic spatial structure, likely due to intraspecific competition, produced a succession of peaks of abundance separated by gaps. At a larger scale (period around 50 m), periodically distributed environmental factors induced aggregates of colonies of species responding positively to these factors. A high quantity of leaf litter and, to a lesser extent, a high density of ground bromeliads promoted a high density and a high species richness of ants. Numerically dominant ants being generally positively associated, interspecific competition was apparently weak. All ant species whose abundance was correlated with an environmental factor were not completely spatially structured by it. This suggests that some other factors, such as intraspecific competition, may have counter-effects. Received 14 March 2005; revised 26 April 2005; accepted 10 May 2005.     

常绿阔叶林林冠环境对栲幼苗建成的影响

植物幼苗建成阶段是决定种群自然更新的关键生活史阶段。研究林冠环境对常绿阔叶林优势种幼苗建成阶段的影响对该类森林的恢复和管理具有重要意义。2014-2016年, 该研究在重庆市缙云山国家级自然保护区的常绿阔叶林的不同林冠环境(大林窗: >150 m ², 中林窗: 100-150 m ², 小林窗: 50-100 m ², 对照: 林下)下进行栲(Castanopsis fargesii)种子野外播种实验, 并对栲幼苗命运和生长情况进行了3年的持续监测。结果表明: (1)栲幼苗出土时间从7月持续到12月, 出苗时间较长, 大林窗对幼苗出土具有延迟作用; (2)栲种子野外平均萌发率为(62.8 ± 2.0)%, 第3个生长季(2016年)末幼苗平均存活率为(65.1 ± 2.2)%, 枯萎是栲幼苗死亡的主要原因; (3)林冠环境对栲种子萌发率及第1个生长季(2014年)末的幼苗存活率无显著影响, 对第2个(2015年)和第3个生长季末的幼苗存活率具有显著影响; (4)林冠环境在第1个生长季对幼苗生长无明显影响, 但在第2个和第3个生长季具有显著影响, 大、中林窗中幼苗总生物量、株高、基径、根长和叶片数显著高于林下, 比叶面积显著低于林下; (5) 3个生长季内, 4类林冠条件下栲幼苗的叶质量比和茎质量比升高, 根质量比和根冠比降低, 并且从第2个生长季开始大林窗中栲幼苗的叶质量比显著高于林下, 根质量比和根冠比显著低于林下。栲幼苗早期的存活和生长依赖种子储存的能量, 受林冠条件影响较弱, 后期则依赖光合作用, 受林冠条件影响较强, 从整个幼苗建成过程看, 大、中林窗更有利于栲幼苗定居。     

干旱与半干旱区灌丛微生境中红砂枯落物分解特征及中小型节肢动物的作用

在干旱区(内蒙古乌拉特后旗)和半干旱区(宁夏盐池)荒漠草原区,选择柠条灌丛内外微生境为研究样地,以红砂枯落物为研究对象,利用2种规格网孔分解袋(30目和250目网孔),探索中小型节肢动物在红砂枯落物分解过程中的作用规律,研究灌丛微生境中红砂枯落物质量损失变化特征及节肢动物群落的贡献。结果表明：(1)在干旱与半干旱区,红砂枯落物分解常数K均表现为灌丛内外微生境间无显著差异,且有无节肢动物参与对K的影响均较小。(2)分解至12个月时,无节肢动物参与的情况下,干旱与半干旱区红砂枯落物残留率均表现为裸地显著低于灌丛;但有节肢动物参与时,枯落物残留率则表现为灌丛内外微生境间无显著差异。分解至44个月时,无节肢动物参与的情况下,仅在干旱区枯落物残留率表现为裸地显著高于灌丛;而有节肢动物参与时,干旱与半干旱区枯落物残留率均表现为灌丛内外微生境间无显著差异。(3)节肢动物对红砂枯落物质量损失的贡献率呈现单峰值现象,且在分解至24个月时达到峰值。枯落物分解至12个月时,仅半干旱区节肢动物对红砂枯落物分解的贡献率表现为裸地显著低于灌丛;分解至44个月时,仅干旱区节肢动物对红砂枯落物分解的贡献率表现为裸地显...     

干旱和半干旱地区灌木下土壤“肥岛”研究进展

世界许多干旱、半干旱地区草地生态系统逐渐被灌木生态系统取代,这种取代过程的发展是由灌木冠幅下土壤中“肥岛”的形成和灌木的扩散相互作用直接造成的,这种“肥岛”的形成和灌木的扩散之间的反馈作用能够改变该系统中的植被组成、结构和土壤养分分布格局,从而改变该生态系统的结构和功能,文中从“肥岛”的概念和形成机制出发,对“肥岛”现象形成的原因、“肥岛”现象的研究意义、研究方法、灌木扩散与“肥岛”之间的相互关系及生物对“肥岛”的响应进行了总结,并分析了“肥岛”研究中应注意的问题,希望为中国干旱、半干旱区域土地退化的成因和过程以及植被恢复的研究提供一些参考。     

Branch direction in Carnegiea gigantea (Cactaceae): Regional patterns and the effect of nurse plants

Abstract. Branch direction in Carnegiea gigantea (saguaro) cacti affects PAR interception and CO₂ uptake, and may influence reception of photomorphogenic cues. Branch direction data were collected in 29 saguaro populations over Arizona to determine if the southerly bias observed in previous studies at two locales can be generalized to the American range of the saguaro, and to determine the effect of shading by nurse plants on branch direction. Locally observed S bias from previous studies can be extended and generalized to include the American saguaro range. Branches were statistically uniform across environmental variability regarding temperature and precipitation. The S bias was not found in saguaros shaded by large nurse plants, supporting the assertion that branch direction occurs, in part, to maximize receipt of insolation.     

The role of cotyledon metabolism in the establishment of quinoa (Chenopodium quinoa) seedlings growing under salinity

A growth chamber experiment was conducted to assess the effect of salinity on emergence, growth, water status, photosynthetic pigments, osmolyte accumulation, and ionic content of quinoa seedlings (Chenopodium quinoa). The aim was to test the hypothesis that quinoa seedlings are well adapted to grow under salinity due to their ability to adjust the metabolic functionality of their cotyledons. Seedlings were grown for 21 days at 250 mM NaCl from the start of the germination. Germination percentage and cotyledon area were not affected by salt whereas seedling height decreased 15%. FW increased in both control and salt-treated cotyledons, but the increase was higher under salinity. DW only increased in salt-treated cotyledons. The DW/FW ratio did not show significant differences between treatments. Relative water content, chlorophyll, carotenoids, lipids, and proteins were significantly lower under salinity. Total soluble sugars, sucrose and glucose concentrations were higher in salt-treated than in control cotyledons. Ion concentration showed a different distribution pattern. Na⁺ and Cl^? concentrations were higher under salinity, while an inverse result was observed for K⁺ concentration. Proline and glycinebetaine concentrations increased under salinity, but the increase was higher in the former than the latter. The osmoprotective role of proline, glycinebetaine, and soluble sugars is discussed.     

Spatial structure and diversity of woody plants and ectomycorrhizal fungus sporocarps in a natural subtropical forest

Spatial patterns of ectomycorrhizal fungi, ectomycorrhizal plants, and non-ectomycorrhizal plants were investigated in a natural subtropical forest using second-order analysis. The results of spatial pattern analysis showed that the degree of clumping of woody plants and ectomycorrhizal sporocarps were correlated. There was a significantly positive correlation of relative aggregation indices between ectomycorrhizal fungi and both non-ectomycorrhizal trees and ectomycorrhizal saplings. Correlations between percentage of ectomycorrhizal trees and sporocarp occurrence of ectomycorrhizal fungi and between diversities of woody plants and ectomycorrhizal fungi were distance-dependent or scale-related. A significantly high percentage of ectomycorrhizal trees was found only at relatively short distance from ectomycorrhizal fungal sporocarps, and significantly positive correlation of the diversity between woody plants and ectomycorrhizal fungi was found only at relative long distance, which implied that ectomycorrhizal sporocarps prefer ectomycorrhizal-tree-dominant micro-sites at near distances and at relatively large scales, diverse ectomycorrhizal sporocarps could be found in woodlands with high diversity of woody plants. Important factors affecting the spatial distribution, occurrence, and diversity of ectomycorrhizal fungi include spatial pattern of ectomycorrhizal plants and non-ectomycorrhizal plants, percentage of ectomycorrhizal plants, and plant diversity in a natural forest.     

The effect of fire and ants on the seed-bank of a shrub in a semi-arid grassland

Abstract Invasion of grasslands by woody plants following the introduction of domestic stock is a worldwide phenomenon. Burning is frequently recommended as a remedial measure but for a pastoral enterprise it is costly and the frequency of the fires required is of critical economic importance. The size and longevity of the soil seed-bank is an essential part of the response of shrub populations to prescribed fire regimes. In this study the seed-bank of the semi-arid zone shrub Dodonaea attenuata in Eragrostis eriopoda tussock grassland was examined in relation to harvesting by ants and the burning history of the sites. On unburnt sites, more than 3500 seeds per m2 entered the seed-bank in the summer 1984–85 but sites burnt 5 years previously produced less than one-third of that number. Burnt shrubs did not flower for 5 years and no seeds survived in the soil through to 1985. Burning immediately prior to seed ripening destroyed the seed crop but burning after seed-fall stimulated a greater germination in the following spring than on unburnt treatments. Ants rapidly harvested most of the seeds produced and after 20 months the combined effects of ant harvesting and germination had reduced the seed-bank at unburnt sites to 8–21 seeds per m2 and at sites burnt 7 years previously to less than two seeds per m2. Seeds were initially harvested in summer by Pheidole spp. of ants for their elaiosome and then discarded in middens outside the entrance to the ant nest. During the subsequent cool season the seeds were taken back into the nest and stored at depths ranging from 2 to 30cm. It was concluded that ants provided short-range dispersal (< 10 m) and promoted the contagious distribution of D. attenuata, which is advantageous for a fire-susceptible, arid-zone shrub invading a grassland liable to be burnt: seedlings derived from seeds in ant storage chambers near the soil surface and in sparse grass situations caused by competition from shrubs may obtain some survival advantage. Prescribed fire has potential as a management tool for controlling population density of D. attenuata because, depending upon season and frequency, it reduces seed rain by killing shrubs, suppresses flowering activity and destroys seed crops on the plant. Under the influence of a regime of regular burning, such as prevailed prior to the European pastoral industry, the limited soil seed-bank would have been a major constraint on D. attenuata populations.     

Spatial variation of the stable nitrogen isotope ratio of woody plants along a topoedaphic gradient in a subtropical savanna

Variation in the stable N isotope ratio (δ¹⁵N) of plants and soils often reflects the influence of environment on the N cycle. We measured leaf δ¹⁵N and N concentration ([N]) on all individuals of Prosopis glandulosa (deciduous tree legume), Condalia hookeri (evergreen shrub), and Zanthoxylum fagara (evergreen shrub) present within a belt transect 308 m long × 12 m wide in a subtropical savanna ecosystem in southern Texas, USA in April and August 2005. Soil texture, gravimetric water content (GWC), total N and δ¹⁵N were also measured along the transect. At the landscape scale, leaf δ¹⁵N was negatively related to elevation for all the three species along this topoedaphic sequence. Changes in soil δ¹⁵N, total N, and GWC appeared to contribute to this spatial pattern of leaf δ¹⁵N. In lower portions of the landscape, greater soil N availability and GWC are associated with relatively high rates of both N mineralization and nitrification. Both soil δ¹⁵N and leaf [N] were positively correlated with leaf δ¹⁵N of non-N₂ fixing plants. Leaf δ¹⁵N of P. glandulosa, an N₂-fixing legume, did not correlate with leaf [N]; the δ¹⁵N of P. glandulosa’s leaves were closer to atmospheric N₂ and significantly lower than those of C. hookeri and Z. fagara. Additionally, at smaller spatial scales, a proximity index (which reflected the density and distance of surrounding P. glandulosa trees) was negatively correlated with leaf δ¹⁵N of C. hookeri and Z. fagara, indicating the N₂-fixing P. glandulosa may be important to the N nutrition of nearby non-N₂-fixing species. Our results indicate plant ¹⁵N natural abundance can reflect the extent of N retention and help us better understand N dynamics and plant-soil interactions at ecosystem and landscape scales.     

The role of the storage carbon of cotyledons in the establishment of seedlings of Hymenaea courbaril under different light conditions

BACKGROUND AND AIMS: Hymenaea courbaril (Leguminosae-Caesalpinioideae) is a tree species with wide distribution through all of the Neotropics. It has large seeds (approx. 5 g) with non-photosynthetic storage cotyledons rich (40 %) in a cell wall polysaccharide (xyloglucan) as a carbon reserve. Because it is found in the understorey of tropical forests, it has been considered as a shade-tolerant, late-secondary species. However, the physiological mechanisms involved in seedling establishment, especially regarding the interplay between storage and light intensity, are not understood. In this work, the ecophysiological role of this carbon cotyledon reserve (xyloglucan) is characterized, emphasizing its effects on seedling growth and development during the transition from heterotrophy to autotrophy under different light conditions. METHODS: Seedlings of H. courbaril were grown in environments with different light intensities, and with or without cotyledons detached before xyloglucan mobilization. Development, growth, photosynthesis and carbon partitioning (dry mass and [14C]sucrose) were analysed in each treatment. KEY RESULTS: The detachment of cotyledons was not important for seedling survival, but resulted in a strong restriction (50 % less) of shoot growth, which was the main sink for the cotyledon carbon reserves. Carbon restriction promoted an early maturation of the photosynthetic apparatus without changes in the net CO2 fixation per unit area. The reduced surface area of the first leaves in seedlings without cotyledons was evidence of limited growth and development of seedlings in low light conditions (22 micromol m(-2) s(-1) photon flux). CONCLUSIONS: There is an increase in the importance of storage xyloglucan in cotyledons for H. courbaril seedling development as light intensity decreases, confirming that this polymer plays a key role in the adaptation of this species to establish successfully in the shadowed understorey of the forest.