Bark in Woody Plants: Understanding the Diversity of a Multifunctional Structure.

Most biological structures carry out multiple functions. Focusing on only one function to make adaptive inferences overlooks that manifold selection pressures and tradeoffs shape the characteristics of a multifunctional structure. Focusing on single functions can only lead to a partial picture of the causes underlying diversity and the evolutionary origin of the structure in question. I illustrate this discussion using bark as a study case. Bark comprises all the tissues surrounding the xylem in woody plants. Broadly, bark includes an inner and mostly living region and an outer, dead one. Of all plant structures, bark has the most complex anatomical structure and ontogenetic origin involving two (and often three) different meristems. Traditionally, the wide diversity in bark traits, mainly bark thickness, has been interpreted as the result of the selective pressures imposed by fire regime. However, recent research has shown that explanations based on fire regime cannot account for salient patterns of bark variation globally including the very strong inner bark thickness-stem diameter scaling, which is likely due to metabolic needs, and the very high intracommunity variation in total, inner, and outer bark thickness, and in inner:outer proportions. Moreover, explanations based on fire disregard that in addition to fire protection, bark carries out several other crucial functions for plants including translocation of photosynthates; storage of starch, soluble sugars, water, and other compounds; protection from herbivores, pathogens, and high temperatures; wound closure, as well as mechanical support, photosynthesis, and likely being involved in xylem embolism repair. All these functions are crucial for plant performance and are involved in synergistic (e.g., storage of water and insulation) and trade-off relationships (e.g., protection from fire vs photosynthetic activity). Focusing on only one of these functions, protection from fire has provided an incomplete picture of the selective forces shaping bark diversity and has severely hindered our incipient understanding of the functional ecology of this crucial region of woody stems. Applying a multifunctional perspective to the study of bark will allow us to address why we observe such high intracommunity variation in bark traits, why some bark trait combinations are ontogenetically impossible or penalized by selection, how bark is coordinated functionally with other plant parts, and as a result, to understand how bark contributes to the vast diversity of plant ecological strategies across the globe.

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