ABSTRACT
To study the acclimation responses of the leaves of saplings of six tree species when changed to low or high levels of irradiance, we carried out a light exposure experiment. Species representative of contrasting shade tolerance groups were identified across a light gradient in the understorey of a Venezuelan Andean cloud forest. Measured traits included gas exchange, chlorophyll fluorescence, and morphoanatomical, biochemical and optical properties. Saplings were grown for 6 months in a shade-house receiving 20% photosynthetic photon flux (PPF) of full sunlight. Plant samples were then moved to shade-houses receiving low PPF (4%) or high PPF (65%). A factorial model (species×PPF), with repeated measurements (0, 15 and 120 days) was designed. Our results showed that morphological and anatomical traits were more plastic to PPF changes than photosynthetic traits. All species were susceptible to photoinhibition (15 days): shade-intolerant species showed dynamic photoinhibition (120 days), whereas shade-tolerant species presented chronic photoinhibition and the consequent inability to increase C assimilation rates under high PPF. The partially shade-tolerant species showed mixed responses; nonetheless, they exhibited larger adjustments in morphoanatomical and optical properties. Thus the acclimation responses of these species when subject to contrasting light conditions could help to explain their distribution along the light gradient in the understorey.
ABSTRACT
The light environment in the understory of cloud forests is highly heterogeneous and determined by species composition, canopy structure, site conditions, and seasonality. This study was carried out at San Eusebio cloud forest, Venezuela (2 300 - 2 500 masl). The impact of canopy structure variations on understory light availability was estimated in the dry (December-February) and rainy (March-November) seasons, in sites under continuous canopy cover and gaps of various sizes. Hemispherical photographs were taken to estimate the percentage of canopy openness, leaf area index, percentages of transmitted direct and diffuse light, and duration and frequency of sunflecks. A light index was calculated from the relative proportions of direct and diffuse light transmitted to the understory. For most variables, there were significant differences between seasons, as well as among different gap sizes and under closed canopy. The light index was low (0.25 to 26 of a maximum = 100), even for the largest gaps, indicating a highly shaded light environment, especially beneath closed canopy in the rainy season. Patterns and interactions among factors were analyzed (gaps vs. continuous canopy, gap sizes, location within the gaps, and seasonality) with a mixed effects repeated measures Anova design. Results showed that the amount of light reaching the understory is low in both gaps and closed canopy. However, small but significant differences in light availability existed for both seasonality and magnitude of the perturbations. These differences could contribute to explain the dynamics of tree species regeneration in this forest. The knowledge of the factors conditioning light availability in the understory where tree regeneration begins is crucial in cloud forests because of energetic limitations in this ecosystem, and might be essential for future restoration and conservation plans concerning the preservation of the diversity and integrity of these forests.
