Optimisation of photosynthetic carbon gain and within-canopy gradients of associated foliar traits for Amazon forest trees
Authors: Lloyd J. , Patiño S. , Paiva R. Q. , Nardoto G. B. , Quesada C. A. , Santos A. J. B. , Baker T. R. , Brand W. A. , Hilke I. , Gielmann H. , Raessler M. , Luizão F. J. , Martinelli L. A. , Mercado L. M. 
Published: June 2010

Abstract. Vertical profiles in leaf mass per unit leaf area (MA), foliar 13C composition (δ13C), nitrogen (N), phosphorus (P), carbon (C) and major cation concentrations were estimated for 204 rain forest trees growing in 57 sites across the Amazon Basin. Data was analysed using a multilevel modelling approach, allowing a separation of gradients within individual tree canopies (within-tree gradients) as opposed to stand level gradients occurring because of systematic differences occurring between different trees of different heights (between-tree gradients). Significant positive within-tree gradients (i.e. increasing values with increasing sampling height) were observed for MA and [C]DW (the subscript denoting on a dry weight basis) with negative within-tree gradients observed for δ13C, [Mg]DW and [K]DW. No significant within-tree gradients were observed for [N]DW, [P]DW or [Ca]DW. The magnitudes of between-tree gradients were not significantly different to the within-tree gradients for MA, δ13C, [C]DW, [K]DW, [N]DW, [P]DW and [Ca]DW. But for [Mg]DW, although there was no systematic difference observed between trees of different heights, strongly negative within-tree gradients were found to occur. When expressed on a leaf area basis (denoted by the subscript "A"), significant positive gradients were observed for [N]A, [P]A and [K]A both within and between trees, these being attributable to the positive intra- and between-tree gradients in MA mentioned above. No systematic within-tree gradient was observed for either [Ca]A or [Mg]A, but with a significant positive gradient observed for [Mg]A between trees (i.e. with taller trees tending to have a higher Mg per unit leaf area). Significant differences in within-tree gradients between individuals were observed only for MA, δ13C and [P] A. This was best associated with the overall average [P]A for each tree, this also being considered to be a surrogate for a tree's average leaf area based photosynthetic capacity, Amax. A new model is presented which is in agreement with the above observations. The model predicts that trees characterised by a low upper canopy Amax should have shallow, or even non-existent, within-canopy gradients in Amax, with optimal intra-canopy gradients becoming sharper as a tree's upper canopy Amax increases. Nevertheless, in all cases it is predicted that the optimal within-canopy gradient in Amax should be substantially less than for photon irradiance. Although this is also shown to be consistent with numerous observations as illustrated by a literature survey of gradients in photosynthetic capacity for broadleaf trees, it is also in contrast to previously held notions of optimality. A new equation relating gradients in photosynthetic capacity within broadleaf tree canopies to the photosynthetic capacity of their upper canopy leaves is presented.