Forest growth is an important factor both economically and ecologically, and it follows a predictable trend with age. Generally, growth accelerates as canopies develop in young forests and declines substantially soon after maximum leaf area is attained. The causes of this decline are multiple and may be linked to age- or size-related processes, or both. Our objective was to determine the relative effects of tree age and tree size on the physiological attributes of two broadleaf species. As age and size are normally coupled during growth, an approach based on grafting techniques to separate the effects of size from those of age was adopted. Genetically identical grafted seedlings were produced from scions taken from trees of four age classes, ranging from 4 to 162 years. We found that leaf-level net photosynthetic rate per unit of leaf mass and some other leaf structural and biochemical characteristics had decreased substantially with increasing size of the donor trees in the field, whereas other gas exchange parameters expressed on a leaf area basis did not. In contrast, these parameters remained almost constant in grafted seedlings, i.e., scions taken from donor trees with different meristematic ages show no age-related trend after they were grafted onto young rootstocks. In general, the results suggested that size-related limitations triggered the declines in photosynthate production and tree growth, whereas less evidence was found to support a role of meristematic age.
Although a substantial body of evidence suggests that large and old trees have reduced metabolic levels, the search for the causes behind this observation has proved elusive. The strong coupling between age and size, commonly encountered in the field, precludes the isolation of the potential causes. We used standard propagation techniques (grafting and air-layering) to decouple the effects of size from those of age in affecting leaf structure, biochemistry and physiology of two broadleaved trees, Acer pseudoplatanus (a diffuse-porous species) and Fraxinus excelsior (a ring-porous species). The first year after establishment of the propagated plants, some of the measurements suggested the presence of age-related declines in metabolism, while other measurements either did not show any difference or suggested variability across treatments not associated with either age or size. During the second year after establishment, only one of the measured properties (specific leaf area) continued to show some evidence of an age-mediated decline (although much reduced compared to the field), whereas, for some properties (particularly for F. excelsior), even the opposite trend of age-related increases was apparent. We concluded that (1) our plants suffered from grafting shock during year 1 and they gradually recovered during year 2; (2) the results over 2 years do not support the statement that age directly mediates ageing in either species but instead suggest that size directly mediates ageing processes; and (3) neither shoots nor roots of A. pseudoplatanus showed any evidence of senescence.
Tree mortality rates appear to be increasing in moist tropical forests (MTFs) with significant carbon cycle consequences. Here, we review the state of knowledge regarding MTF tree mortality, create a conceptual framework with testable hypotheses regarding the drivers, mechanisms and interactions that may underlie increasing MTF mortality rates, and identify the next steps for improved understanding and reduced prediction. Increasing mortality rates are associated with rising temperature and vapor pressure deficit, liana abundance, drought, wind events, fire and, possibly, CO2 fertilization-induced increases in stand thinning or acceleration of trees reaching larger, more vulnerable heights. The majority of these mortality drivers may kill trees in part through carbon starvation and hydraulic failure. The relative importance of each driver is unknown. High species diversity may buffer MTFs against large-scale mortality events, but recent and expected trends in mortality drivers give reason for concern regarding increasing mortality within MTFs. Models of tropical tree mortality are advancing the representation of hydraulics, carbon and demography, but require more empirical knowledge regarding the most common drivers and their subsequent mechanisms. We outline critical datasets and model developments required to test hypotheses regarding the underlying causes of increasing MTF mortality rates, and improve prediction of future mortality under climate change.