Contents
 	Summary	30	
I.	Allocation in perspective	31	
II.	Topics of this review	32	
III.	Methodology	32	
IV.	Environmental effects	33	
V.	Ontogeny	36	
VI.	Differences between species	40	
VII.	Physiology and molecular regulation	41	
VIII.	Ecological aspects	42	
IX.	Perspectives	45	
 	Acknowledgements	45	
 	References	45	
 	Appendices A1–A4	49	


Summary
We quantified the biomass allocation patterns to leaves, stems and roots in vegetative plants, and how this is influenced by the growth environment, plant size, evolutionary history and competition. Dose–response curves of allocation were constructed by means of a meta-analysis from a wide array of experimental data. They show that the fraction of whole-plant mass represented by leaves (LMF) increases most strongly with nutrients and decreases most strongly with light. Correction for size-induced allocation patterns diminishes the LMF-response to light, but makes the effect of temperature on LMF more apparent. There is a clear phylogenetic effect on allocation, as eudicots invest relatively more than monocots in leaves, as do gymnosperms compared with woody angiosperms. Plants grown at high densities show a clear increase in the stem fraction. However, in most comparisons across species groups or environmental factors, the variation in LMF is smaller than the variation in one of the other components of the growth analysis equation: the leaf area : leaf mass ratio (SLA). In competitive situations, the stem mass fraction increases to a smaller extent than the specific stem length (stem length : stem mass). Thus, we conclude that plants generally are less able to adjust allocation than to alter organ morphology.

https://nph.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1469-8137.2011.03952.x

Biomass allocation to leaves, stems and roots: meta-analyses of interspecific variation and environmental control

1. Water availability is the main determinant of species' distribution in lowland tropical forests. Species' occurrence along water availability gradients depends on their ability to tolerate drought. 2. To identify species' traits underlying drought-tolerance we excavated first year seedlings of 62 dry and moist forest tree species at the onset of the dry season. We evaluate how morphological seedling traits differ between forests, and whether functional groups of species can be identified based on trait relations. We also compare seedling traits along independent axes of drought and shade-tolerance to assess a hypothesized trade-off. 3. Seedlings of dry forest species improve water foraging capacity in deep soil layers by an increased below-ground biomass allocation and by having deep roots. They minimize the risk of cavitation by making dense stems, and reduce transpiration by producing less leaf tissue. Moist forest seedlings have large leaf areas and a greater above-ground biomass, to maximize light interception, and long, cheap, branched root systems, to increase water and nutrient capture. 4. Associations among seedling traits reveal three major drought strategies: (i) evergreen drought-tolerant species have high biomass investment in enduring organs, minimize cavitation and minimize transpiration to persist under dry conditions; (ii) drought-avoiding species maximize resource capture during a limited growing season and then avoid stress with a deciduous leaf habit in the dry season; (iii) drought-intolerant species maximize both below- and above-ground resource capture to increase competitiveness for light, but are consequently precluded from dry habitats. 5. We found no direct trade-off between drought- and shade-tolerance, because they depend largely on different morphological adaptations. Drought-tolerance is supported by a high biomass investment to the root system, whereas shade-tolerance is mainly promoted by a low growth rate and low SLA. 6. Synthesis. We conclude that there are three general adaptation strategies of drought-tolerance, which seemingly hold true across biomes and for different life forms. Drought- and shade-tolerance are largely independent from one another, suggesting a high potential for niche differentiation, as species' specialization can occur at different combinations of water and light availability.

https://besjournals.onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-2745.2008.01466.x

Seedling root morphology and biomass allocation of 62 tropical tree species in relation to drought- and shade-tolerance

Global environmental change, related to climate change and the deposition of airborne N-containing contaminants, has already resulted in shifts in plant community composition among plant functional types in Arctic and temperate alpine regions. In this paper, we review how key ecosystem processes will be altered by these transformations, the complex biological cascades and feedbacks that might result, and some of the potential broader consequences for the earth system. Firstly, we consider how patterns of growth and allocation, and nutrient uptake, will be altered by the shifts in plant dominance. The ways in which these changes may disproportionately affect the consumer communities, and rates of decomposition, are then discussed. We show that the occurrence of a broad spectrum of plant growth forms in these regions (from cryptogams to deciduous and evergreen dwarf shrubs, graminoids and forbs), together with hypothesized low functional redundancy, will mean that shifts in plant dominance result in a complex series of biotic cascades, couplings and feedbacks which are supplemental to the direct responses of ecosystem components to the primary global change drivers. The nature of these complex interactions is highlighted using the example of the climate-driven increase in shrub cover in low-Arctic tundra, and the contrasting transformations in plant functional composition in mid-latitude alpine systems. Finally, the potential effects of the transformations on ecosystem properties and processes that link with the earth system are reviewed. We conclude that the effects of global change on these ecosystems, and potential climate-change feedbacks, cannot be predicted from simple empirical relationships between processes and driving variables. Rather, the effects of changes in species distributions and dominances on key ecosystem processes and properties must also be considered, based upon best estimates of the trajectories of key transformations, their magnitude and rates of change.

/pdf/ecosystem-feedbacks-and-cascade-processes-understanding-5dedw57w3u.pdf

Ecosystem feedbacks and cascade processes: understanding their role in the responses of Arctic and alpine ecosystems to environmental change

Summary
1Optimal partitioning theory (OPT) suggests that plants should allocate biomass to the organ that acquires the most limiting resource. An implied assumption of this is that there are trade-offs in allocation between leaf, stem and root functions.
2Recently, an alternative approach, allometric biomass partitioning theory (APT), was developed to predict how plants should divide their metabolic production between leaves, stems and roots, based on the constraints of body size. APT predicts that, for an allometrically ideal plant, leaf mass should scale to the 3/4th power of body size, and stem and root mass should scale isometrically to body size.
3In this study, we combine OPT with APT by investigating biomass partitioning not accounted for by allometric constraints across broad environmental gradients.
4Intraspecific variability in biomass allocation shows correlations with environmental factors that would be predicted by OPT. However, interspecifically, these patterns either do not appear or are greatly reduced.
5Our study suggests that, after size is accounted for, intraspecific residual variation in biomass allocation may be partially explained by environmental factors in a manner consistent with OPT. However, the particular patterns vary between species, obscuring large-scale patterns.
6In summary, differences due to environmental variability can be incorporated with allocation patterns related to total mass to help understand how plants should allocate biomass in response to changes in both size and environment.

https://besjournals.onlinelibrary.wiley.com/doi/epdf/10.1111/j.1365-2435.2007.01276.x

Consistency between an allometric approach and optimal partitioning theory in global patterns of plant biomass allocation

Leaf phenology remains one of the most difficult processes to parameterize in terrestrial ecosystem models because our understanding of the physical processes that initiate leaf onset and senescence is incomplete. While progress has been made at the molecular level, for example by identifying genes that are associated with senescence and flowering for selected plant species, a picture of the processes controlling leaf phenology is only beginning to emerge. A variety of empirical formulations have been used with varying degrees of success in terrestrial ecosystem models for both extratropical and tropical biomes. For instance, the use of growing degree-days (GDDs) to initiate leaf onset has received considerable recognition and this approach is used in a number of models. There are, however, limitations when using GDDs and other empirically based formulations in global transient climate change simulations. The phenology scheme developed for the Canadian Terrestrial Ecosystem Model (CTEM), designed for inclusion in the Canadian Centre for Climate Modelling and Analysis coupled general circulation model, is described. The representation of leaf phenology is general enough to be applied over the globe and sufficiently robust for use in transient climate change simulations. Leaf phenology is functionally related to the (possibly changing) climate state and to atmospheric composition rather than to geographical boundaries or controls implicitly based on current climate. In this approach, phenology is controlled by environmental conditions as they affect the carbon balance. A carbon-gain-based scheme initiates leaf onset when it is beneficial for the plant, in carbon terms, to produce new leaves. Leaf offset is initiated by unfavourable environmental conditions that incur carbon losses and these include shorter day length, cooler temperatures, and dry soil moisture conditions. The comparison of simulated leaf onset and offset times with observation-based estimates for temperate and boreal deciduous, tropical evergreen, and tropical deciduous plant functional types at selected locations indicates that the phenology scheme performs satisfactorily. Model simulated leaf area index and stem and root biomass are also compared with observational estimates to illustrate the performance of CTEM.

A parameterization of leaf phenology for the terrestrial ecosystem component of climate models

Relative growth rate (RGR) and other growth parameters were studied in eight pairs of closely related deciduous and evergreen species (within the same genus or family). The main objective of this study was to test the association between leaf turnover rate and RGR, specific leaf area (SLA, leaf area/leaf dry weight) and other growth variables. Plants were grown for 6 months in a greenhouse under favourable water and nutrient conditions. Variation in RGR among the 16 woody species was due mainly to differences in morphological parameters such as leaf area ratio (LAR, whole plant area/whole plant dry weight) and SLA). However, temporal variation in RGR within species was due mainly to variation in net assimilation rate. When phylogeny was not taken into account, analyses showed that deciduous species grew faster than evergreens. In contrast, when phylogeny was taken into account, the data analysis showed that a faster RGR is not consistently associated with the deciduous habit (in five pairs it was, but in the other three it was not). The faster growth of the deciduous trees (in the five positive contrasts) could be explained by their higher LAR and higher SLA relative to evergreens. The lack of differences in RGR between deciduous and evergreens (in three pairs) was due to the higher leaf mass ratio (LMR, leaf dry biomass/total dry biomass) for the evergreens, which offset the higher SLA of the deciduous species, resulting in a similar LAR in both functional groups (LAR=LMR×SLA). Deciduous species had consistently higher SLA than evergreens. We suggest that SLA, more than RGR, could be an important parameter in determining adaptive advantages of deciduous and evergreen species.

/pdf/relative-growth-rate-in-phylogenetically-related-deciduous-35hrfq4hp0.pdf

Isabel Antúnez

Papers

Relative growth rate in phylogenetically related deciduous and evergreen woody species