RESUMO
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.
RESUMO
Resumen:El ambiente lumínico en el sotobosque de las selvas nubladas es muy heterogéneo y está determinado por la composición de especies, la estructura del dosel, las condiciones de sitio y la estacionalidad. Este estudio se realizó en la selva nublada San Eusebio, Venezuela (2 300-2 500 m.s.n.m.). El impacto de las variaciones estructurales del dosel en la disponibilidad de luz del sotobosque fue estimado durante la estación seca (diciembre-febrero) y lluviosa (marzo-noviembre), tanto en claros de diferente tamaño como en bosque no perturbado (BNP). Se tomaron fotografías hemisféricas para estimar el porcentaje de dosel abierto, índice de área foliar, porcentajes de luz directa y luz difusa transmitida, duración y frecuencia de rayos de sol. Se calculó un índice de luz a partir de las proporciones relativas de luz directa y difusa transmitida al sotobosque. Para la mayoría de las variables, hubo diferencias significativas entre las estaciones, así como entre claros y BNP. El índice de luz fue bajo (0.25 a 26 de un máximo = 100), incluso para los claros más grandes, indicando un ambiente de luz muy oscuro en el sotobosque, particularmente, en BNP en la estación lluviosa. Patrones e interacciones entre los factores fueron analizados (BNP vs. claros, tamaño de claro, posición en el claro y estacionalidad) con un diseño ANOVA de efectos mixtos y medidas repetidas. Los resultados mostraron que la cantidad de luz que llega al sotobosque es baja, tanto en BNP como en claros. Sin embargo, existieron diferencias pequeñas pero significativas en la disponibilidad de luz, tanto por efecto de la estacionalidad como por la magnitud de las perturbaciones. Estas diferencias podrían contribuir a explicar la dinámica de la regeneración de especies en estos bosques. El conocimiento de los factores que condicionan la disponibilidad de luz en el sotobosque, donde se inicia la regeneración de los árboles, es crucial en selvas nubladas debido a las limitaciones energéticas de este ecosistema, y podría ser fundamental para futuros planes de restauración y conservación de la diversidad e integridad de estos bosques.
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. Rev. Biol. Trop. 64 (4): 1699-1707. Epub 2016 December 01.
Assuntos
Estações do Ano , Luz Solar , Florestas , Clima , Chuva , Valores de Referência , Fatores de Tempo , Venezuela , BiodiversidadeRESUMO
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.
Assuntos
Clima , Florestas , Estações do Ano , Luz Solar , Biodiversidade , Chuva , Valores de Referência , Fatores de Tempo , VenezuelaRESUMO
Chamaedorea bartlingiana is a dioecious palm that grows in the cloud forest understories of the Venezuelan Andes. Age and sexual differences in phenology and reproductive patterns were studied in labelled individuals of all age categories. This species has long-lived leaves and low leaf production, both characteristic of understory plants. Growth rates are lower in juveniles than in adults and in females than in males, as in other palms. Male and female individuals show different reproductive patterns. Male inflorescences are always produced at the same rate and the probability of surviving until anthesis is constant. Females produce reproductive buds at the same rate as males, but these buds have a 35% probability of becoming a ripe infrutescence if the plant has infrutescences already growing, and 70% if it does not. This pattern and the slow growth of inflorescences (1 year for males from bud to flowers, 2 years for females from bud to ripe fruits) cause a pluriannual reproductive pattern at the population level. Field germination does not follow this pattern, but shows one annual peak probably related to environmental conditions.