Tuesday, October 27, 2009

When will SLA R.I.P.?

Relationship between leaf tissue density and the abundance of grassland species in uplands at Konza Prairie. Each point is a different species with its abundance measured over 14 years.

For almost two decades, SLA (or its inverse alter-ego, LMA) has reigned supreme as the central functional trait of plants. SLA, i.e. specific leaf area--the ratio of leaf area to mass, has stood to represent the amount of investment into light acquisition. Entire pyramids of approaches to traits are built on the fundamental supremacy of SLA. The only thing more important than SLA in these pyramids is relative growth rate (RGR).

But why SLA? Why the ratio of area to mass? The thinking is that plants that grow fast need to absorb as much light as possible with the least amount of investment. Hence, selection favors plants that produce a lot of leaf area with little carbon investment, i.e. a high SLA. Plants in stressful or low-resource areas have low SLA, which presumably aids plants in resisting stress or maximizing the utilization of a limiting resource. Consistently, there are good correlations between SLA and RGR as well as other leaf characteristics such as photosynthetic rates, which have reinforced the primacy of SLA.

For almost all of the 20 years, there has always been a countervailing opinion of SLA that has never been rectified. If it ever was squared, SLA would likely never be measured again.

A leaf can high SLA either because it is thin or because it has low tissue density—thickness and density are the two components of SLA. In 1991, Witkowski and Lamont examined thickness and density across a series of ecological contrasts for sclerophyllous species. In short, from the patterns they observed, the authors concluded that “leaf density and thickness may respond to independently to resource and other gradients, and thus are more appropriate measures than [SLA] which confounds them.” Because thickness is so easy to measure—a quick squeezing of calipers—there is no good reason to not break down SLA to density and thickness every time.

Thickness and density have different functional roles in a leaf. They often vary independently across ecological contrasts. A thick, low density leaf and a thin, high density leaf would have the same SLA, but very different performances in most environments. By extension, SLA might be important to plant ecologists, but not to selection.

But maybe this is a bit hasty. SLA is supposed to be ecologically important and help explain the abundance of species across contrasts. Maybe SLA explains abundance better than thickness or tissue density. Surprisingly, the relative explanatory of SLA and its components have rarely been tested quantitatively. In general, this is probably the Achilles heel of most traits work. We spend more time examining relationships among traits than rigorously testing their relative predictive capacity.

Refuting the ecological importance of SLA or either of its components will not be a simple affair. It’ll take a number of studies before we understand their relative empirical importance. I’ve now done two. The first was at Cedar Creek along fertilization and disturbance gradients. The second is at Konza where I measured leaf traits for 130 grassland herbaceous species and tested their predictive capacity for species abundance across topographic, burning, and grazing contrasts. The results for Konza? SLA explained no variation in the abundance of species. Yet, tissue density did. Consistently across gradients it was tissue density not SLA that explained the abundance of species. The Cedar Creek work largely concluded the same thing.

SLA should not be buried yet, but at some point, we are going to have to fundamentally reexamine the hierarchy of traits in the ecology of plants. A dichotomous world of high SLA and low SLA (if not high RGR and low RGR) plant species might have to be replaced. Until then, at the very least, measure thickness.

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