RESUMEN
Increases in hydrological extremes, including drought, are expected for Amazon forests. A fundamental challenge for predicting forest responses lies in identifying ecological strategies which underlie such responses. Characterization of species-specific hydraulic strategies for regulating water-use, thought to be arrayed along an 'isohydric-anisohydric' spectrum, is a widely used approach. However, recent studies have questioned the usefulness of this classification scheme, because its metrics are strongly influenced by environments, and hence can lead to divergent classifications even within the same species. Here, we propose an alternative approach positing that individual hydraulic regulation strategies emerge from the interaction of environments with traits. Specifically, we hypothesize that the vertical forest profile represents a key gradient in drought-related environments (atmospheric vapor pressure deficit, soil water availability) that drives divergent tree water-use strategies for coordinated regulation of stomatal conductance (gs) and leaf water potentials (ΨL) with tree rooting depth, a proxy for water availability. Testing this hypothesis in a seasonal eastern Amazon forest in Brazil, we found that hydraulic strategies indeed depend on height-associated environments. Upper canopy trees, experiencing high vapor pressure deficit (VPD), but stable soil water access through deep rooting, exhibited isohydric strategies, defined by little seasonal change in the diurnal pattern of gs and steady seasonal minimum ΨL. In contrast, understory trees, exposed to less variable VPD but highly variable soil water availability, exhibited anisohydric strategies, with fluctuations in diurnal gs that increased in the dry season along with increasing variation in ΨL. Our finding that canopy height structures the coordination between drought-related environmental stressors and hydraulic traits provides a basis for preserving the applicability of the isohydric-to-anisohydric spectrum, which we show here may consistently emerge from environmental context. Our work highlights the importance of understanding how environmental heterogeneity structures forest responses to climate change, providing a mechanistic basis for improving models of tropical ecosystems.
Asunto(s)
Bosques , Árboles , Agua , Agua/metabolismo , Agua/fisiología , Árboles/fisiología , Brasil , Sequías , Transpiración de Plantas/fisiología , Suelo/química , Hojas de la Planta/fisiologíaRESUMEN
Plant phenology-the timing of cyclic or recurrent biological events in plants-offers insight into the ecology, evolution, and seasonality of plant-mediated ecosystem processes. Traditionally studied phenologies are readily apparent, such as flowering events, germination timing, and season-initiating budbreak. However, a broad range of phenologies that are fundamental to the ecology and evolution of plants, and to global biogeochemical cycles and climate change predictions, have been neglected because they are "cryptic"-that is, hidden from view (e.g., root production) or difficult to distinguish and interpret based on common measurements at typical scales of examination (e.g., leaf turnover in evergreen forests). We illustrate how capturing cryptic phenology can advance scientific understanding with two case studies: wood phenology in a deciduous forest of the northeastern USA and leaf phenology in tropical evergreen forests of Amazonia. Drawing on these case studies and other literature, we argue that conceptualizing and characterizing cryptic plant phenology is needed for understanding and accurate prediction at many scales from organisms to ecosystems. We recommend avenues of empirical and modeling research to accelerate discovery of cryptic phenological patterns, to understand their causes and consequences, and to represent these processes in terrestrial biosphere models.
Asunto(s)
Ecosistema , Bosques , Brasil , Cambio Climático , Estaciones del AñoRESUMEN
Satellite and tower-based metrics of forest-scale photosynthesis generally increase with dry season progression across central Amazônia, but the underlying mechanisms lack consensus. We conducted demographic surveys of leaf age composition, and measured the age dependence of leaf physiology in broadleaf canopy trees of abundant species at a central eastern Amazon site. Using a novel leaf-to-branch scaling approach, we used these data to independently test the much-debated hypothesis - arising from satellite and tower-based observations - that leaf phenology could explain the forest-scale pattern of dry season photosynthesis. Stomatal conductance and biochemical parameters of photosynthesis were higher for recently mature leaves than for old leaves. Most branches had multiple leaf age categories simultaneously present, and the number of recently mature leaves increased as the dry season progressed because old leaves were exchanged for new leaves. These findings provide the first direct field evidence that branch-scale photosynthetic capacity increases during the dry season, with a magnitude consistent with increases in ecosystem-scale photosynthetic capacity derived from flux towers. Interactions between leaf age-dependent physiology and shifting leaf age-demographic composition are sufficient to explain the dry season photosynthetic capacity pattern at this site, and should be considered in vegetation models of tropical evergreen forests.
Asunto(s)
Carbono/metabolismo , Bosques , Hojas de la Planta/fisiología , Estaciones del Año , Brasil , Clorofila/metabolismo , Gases/metabolismo , Fotosíntesis , Estomas de Plantas/fisiología , Factores de TiempoRESUMEN
Leaf age structures the phenology and development of plants, as well as the evolution of leaf traits over life histories. However, a general method for efficiently estimating leaf age across forests and canopy environments is lacking. Here, we explored the potential for a statistical model, previously developed for Peruvian sunlit leaves, to consistently predict leaf ages from leaf reflectance spectra across two contrasting forests in Peru and Brazil and across diverse canopy environments. The model performed well for independent Brazilian sunlit and shade canopy leaves (R2 = 0.75-0.78), suggesting that canopy leaves (and their associated spectra) follow constrained developmental trajectories even in contrasting forests. The model did not perform as well for mid-canopy and understory leaves (R2 = 0.27-0.29), because leaves in different environments have distinct traits and trait developmental trajectories. When we accounted for distinct environment-trait linkages - either by explicitly including traits and environments in the model, or, even better, by re-parameterizing the spectra-only model to implicitly capture distinct trait-trajectories in different environments - we achieved a more general model that well-predicted leaf age across forests and environments (R2 = 0.79). Fundamental rules, linked to leaf environments, constrain the development of leaf traits and allow for general prediction of leaf age from spectra across species, sites and canopy environments.