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          <family>Lindh</family>
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    <title>Constrained growth flips the direction of optimal phenological responses among annual plants</title>
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    <keywords>climate change; constrained growth; flowering; history; optimal control theory; phenology; productivity; season</keywords>
    <abstract>Phenological changes among plants due to climate change are well documented, but often hard to interpet. In order to assess the adaptive value of observed changes, we study how annual plants with and without growth constraints should optimize their flowering time when productivity and season length changes. We consider growth constraints that depend on the plant&apos;s  vegetative mass: self-shading, costs for nonphotosynthetic structural tissue and sibling competition. 



We derive the optimal flowering time from a dynamic energy allocation model using optimal control theroy. We prove that an  immediate switch (bang-bang control) from vegetative to reproductive growth is optimal with constrained growth and constant mortality. 



Increasing mean productivity, while keeping season length constant and growth unconstrained, delayed the optimal flowering  time. When growth was constrained and productivity was relatively high, the optimal flowering time advanced instead. When the growth season was extended equally at both ends, the optimal flowering time was advanced under constrined growth and delayed under unconstrained growth. 



Our results suggests that growth constraints are key factors to consider when interpreting phenologicl flowering responses. It  can help to explain phenological patterns along productivity gradients, and links empirical observations made on calendar scales with life-history theory.</abstract>
    <date>2016-03</date>
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    <iiasa_bibref>New Phytologist (Published online 9 November 2015)</iiasa_bibref>
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