Phylogenetic analysis reveals a scattered distribution of autumn colours

Background and Aims

Leaf colour in autumn is rarely considered informative for taxonomy, but there is now growing interest in the evolution of autumn colours and different hypotheses are debated. Research efforts are hindered by the lack of basic information: the phylogenetic distribution of autumn colours. It is not known when and how autumn colours evolved.

Methods

Data are reported on the autumn colours of 2368 tree species belonging to 400 genera of the temperate regions of the world, and an analysis is made of their phylogenetic relationships in order to reconstruct the evolutionary origin of red and yellow in autumn leaves.

Key Results

Red autumn colours are present in at least 290 species (70 genera), and evolved independently at least 25 times. Yellow is present independently from red in at least 378 species (97 genera) and evolved at least 28 times.

Conclusions

The phylogenetic reconstruction suggests that autumn colours have been acquired and lost many times during evolution. This scattered distribution could be explained by hypotheses involving some kind of coevolutionary interaction or by hypotheses that rely on the need for photoprotection.Key words: Autumn colour, leaf colour, comparative analysis, coevolution, photoprotection, phylogenetic analysis     

Demographic consequences of chromatic leaf defence in tropical tree communities: do red young leaves increase growth and survival?

Background

Many tropical forest tree species delay greening their leaves until full expansion. This strategy is thought to provide newly flushing leaves with protection against damage by herbivores by keeping young leaves devoid of nutritive value. Because young leaves suffer the greatest predation from invertebrate herbivores, delayed greening could prevent costly tissue loss. Many species that delay greening also produce anthocyanin pigments in their new leaves, giving them a reddish tint. These anthocyanins may be fungicidal, protect leaves against UV damage or make leaves cryptic to herbivores blind to the red part of the spectrum.

Methods

A comprehensive survey was undertaken of seedlings, saplings and mature trees in two diverse tropical forests: a rain forest in western Amazonia (Yasuní National Park, Ecuador) and a deciduous forest in Central America (Barro Colorado Island, Panamá). A test was made of whether individuals and species with delayed greening or red-coloured young leaves showed lower mortality or higher relative growth rates than species that did not.

Key results

At both Yasuní and Barro Colorado Island, species with delayed greening or red young leaves comprised significant proportions of the seedling and tree communities. At both sites, significantly lower mortality was found in seedlings and trees with delayed greening and red-coloured young leaves. While there was little effect of leaf colour on the production of new leaves of seedlings, diameter relative growth rates of small trees were lower in species with delayed greening and red-coloured young leaves than in species with regular green leaves, and this effect remained when the trade-off between mortality and growth was accounted for.

Conclusions

Herbivores exert strong selection pressure on seedlings for the expression of defence traits. A delayed greening or red-coloured young leaf strategy in seedlings appears to be associated with higher survival for a given growth rate, and may thus influence the species composition of later life stages.     

The role of forest tent caterpillar defoliations and partial harvest in the decline and death of sugar maple

Background and Aims

Natural and anthropogenic disturbances can act as stresses on tree vigour. According to Manion''s conceptual model of tree disease, the initial vigour of trees decreases as a result of predisposing factors that render these trees more vulnerable to severe inciting stresses, stresses that can then cause final vigour decline and subsequent tree death. This tree disease model was tested in sugar maple (Acer saccharum) by assessing the roles of natural and anthropogenic disturbances in tree decline and death.

Methods

Radial growth data from 377 sugar maple trees that had undergone both defoliations by insects and partial harvest were used to estimate longitudinal survival probabilities as a proxy for tree vigour. Radial growth rates and survival probabilities were compared among trees subjected to different levels of above- and below-ground disturbances, between periods of defoliation and harvest, and between live and dead trees.

Key Results

Manion''s tree disease model correctly accounts for vigour decline and tree death in sugar maple; tree growth and vigour were negatively affected by a first defoliation, predisposing these trees to death later during the study period due to a second insect outbreak that initiated a final vigour decline. This decline was accelerated by the partial harvest disturbance in 1993. Even the most severe anthropogenic disturbances from partial harvest did not cause, unlike insect defoliation, any growth or vigour declines in live sugar maple.

Conclusions

Natural disturbances acted as predisposing and inciting stresses in tree sugar maple decline and death. Anthropogenic disturbances from a partial harvest at worst accelerated a decline in trees that were already weakened by predisposing and inciting stresses (i.e. repeated insect defoliations). Favourable climatic conditions just before and after the partial harvest may have alleviated possible negative effects on growth resulting from harvesting.Key words: Manion, tree disease model, disturbance, Acer saccharum, tree mortality, tree vigour     

A field study with geometrid moths to test the coevolution hypothesis of red autumn colours in deciduous trees

Red autumn colouration of trees is the result of newly synthesized anthocyanin pigments in senescing autumn leaves. As anthocyanin accumulation is costly and the trait is not present in all species, anthocyanins must have an adaptive significance in autumn leaves. According to the coevolution hypothesis of autumn colours, red autumn leaves warn herbivorous insects – especially aphids that migrate to reproduce in trees in the autumn – that the tree will not be a suitable host for their offspring in spring due to a high level of chemical defence or lack of nutrients. The signalling allows trees to avoid herbivores and herbivores to choose better host trees. In this study the coevolution hypothesis was tested with four deciduous tree species that have red autumn leaf colouration – European aspen (Populus tremula L.) (Salicaceae), rowan (Sorbus aucuparia L.) (Rosaceae), mountain birch [Betula pubescens ssp. czerepanovii (NI Orlova) Hämet‐Ahti], and dwarf birch (Betula nana L.) (Betulaceae), and with two generalist herbivores, the autumnal moth [Epirrita autumnata (Borkhausen)] and the winter moth [Operophtera brumata (L.)] (both Lepidoptera: Geometridae). Anthocyanin concentrations of autumn leaves were determined from leaf samples and the growth performance parameters of the moth larvae on the study trees were measured in the spring. Trees with higher anthocyanin concentration in the autumn were predicted to be low‐quality food for the herbivores. Our results clearly showed that anthocyanin concentration was not correlated with the growth performance of the moths in any of the studied tree species. Thus, our study does not support the coevolution hypothesis of autumn colours.     

Spring versus autumn leaf colours: Evidence for different selective agents and evolution in various species and floras

Red colouration is common in young and old leaves of broadleaf woody species. Assuming that leaf colours are adaptive, we examined, by comparing the colouration in young versus old leaves, the possibility that different selection agents may have operated on spring versus autumn leaf colouration. We observed spring versus autumn colouration in three very different woody floras (Finland, Japan and Israel) in order to allow for a broad ecological and evolutionary spectrum. The null hypothesis was that if the same selective agents operated in spring and autumn, it is expected that when spring leaves are red, they should always be red in autumn, and when spring leaves are green, they should be green or yellow in autumn. We found that green spring leaves are almost exclusively associated with yellow leaf colour at senescence in autumn. Species with red autumn leaves almost always have at least some red colouration in their spring leaves. However, about half of the species with red spring leaves have yellow autumn leaves. Brown autumn leaves were not common in the species we studied. As about half of the species with red spring leaves have yellow autumn leaves but not vice versa, we conclude that there are many cases in which the selecting agents for spring versus autumn leaf colour were not the same.     

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Evidence from the domestication of apple for the maintenance of autumn colours by coevolution

The adaptive value of autumn colours is still a puzzle for evolutionary biology. It has been suggested that autumn colours are a warning signal to insects that use the trees as a host. I show that aphids (Dysaphis plantaginea) avoid apple trees (Malus pumila) with red leaves in autumn and that their fitness in spring is lower on these trees, which suggests that red leaves are an honest signal of the quality of the tree as a host. Autumn colours are common in wild populations but not among cultivated apple varieties, which are no longer under natural selection against insects. I show that autumn colours remain only in the varieties that are very susceptible to the effects of a common insect-borne disease, fire blight, and therefore are more in need of avoiding insects. Moreover, varieties with red leaves have smaller fruits, which shows that they have been under less effective artificial selection. This suggests a possible trade off between fruit size, leaf colour and resistance to parasites. These results are consistent with the hypothesis that autumn colours are a warning signal to insects, but not with other hypotheses.     

Testing the enemy release hypothesis: a comparison of foliar insect herbivory of the exotic Norway maple (Acer platanoides L.) and the native sugar maple (A. saccharum L.)

Norway maple (Acer platanoides) is a Eurasian introduced tree species which has invaded the North American range of its native congener, sugar maple (A. saccharum). One hypothesis used to explain the success of an invasive species is the enemy release hypothesis (ERH), which states that invasive species are often particularly successful in their new range because they lack the enemies of their native range. In this study, we hypothesized that Norway maple would have less insect damage than sugar maple due to such enemy release. Autumn 2005 and summer 2006 leaves of Norway and sugar maple were collected from six sites in New Jersey and Pennsylvania to compare percent leaf area loss, gall damage, fungal damage, and specific leaf area (cm²/g). Although both species had low overall mean levels of leaf damage (0.4–2.5%), in both years/seasons Norway maple had significantly less leaf damage than sugar maple. Insects were also collected to compare insect assemblies present on each tree species. The numbers of insect taxa and individuals found on each species were nearly equivalent. Overall, the results of this study are consistent with the enemy release hypothesis for Norway maple. In addition, sugar maples when surrounded by Norway maples tended to show reduced herbivory. This suggests that the spread of Norway maple in North America, by reducing amounts of insect herbivory, may have further ecosystem-wide impacts.     

Importance of olfactory and visual signals of autumn leaves in the coevolution of aphids and trees

Deciduous trees remobilize the nitrogen in senescing leaves during the process of autumn colouration, which in many species is associated with increased concentrations of anthocyanins. Archetti and Hamilton and Brown observed that autumn colouration is stronger in tree species facing a high diversity of specialist aphids. They proposed a coevolution theory that the bright colours in autumn might provide an honest signal of defence commitment, thus deterring migrant aphids from settling on the leaves. So far, there have been very few experimental results to support the hypothesis, and tree commitment to phenolics-based defences has not shown direct protection against aphids. Predators and parasitoids have been found to be the major controllers of arboreal aphids. Indirect defences involve the emission of attractive volatile compounds that enhance the effectiveness of carnivorous enemies. The indirect defence hypothesis is presented to explain low aphid diversity on tree species that are green during autumn. The hypothesis suggests that green foliage can continue to produce herbivore-inducible plant volatiles and maintain volatile-based indirect plant defences against aphids until leaf abscission.     

Common allometric response of open-grown leader shoots to tree height in co-occurring deciduous broadleaved trees

Background and Aims

Morphology of crown shoots changes with tree height. The height of forest trees is usually correlated with the light environment and this makes it difficult to separate the effects of tree size and of light conditions on the morphological plasticity of crown shoots. This paper addresses the tree-height dependence of shoot traits under full-light conditions where a tree crown is not shaded by other crowns.

Methods

Focus is given to relationships between tree height and top-shoot traits, which include the shoot''s leaf-blades and non-leafy mass, its total leaf-blade area and the length and basal diameter of the shoot''s stem. We examine the allometric characteristics of open-grown current-year leader shoots at the tops of forest tree crowns up to 24 m high and quantify their responses to tree height in 13 co-occurring deciduous hardwood species in a cool-temperate forest in northern Japan.

Key Results

Dry mass allocated to leaf blades in a leader shoot increased with tree height in all 13 species. Specific leaf area decreased with tree height. Stem basal area was almost proportional to total leaf area in a leader shoot, where the proportionality constant did not depend on tree height, irrespective of species. Stem length for a given stem diameter decreased with tree height.

Conclusions

In the 13 species observed, height-dependent changes in allometry of leader shoots were convergent. This finding suggests that there is a common functional constraint in tree-height development. Under full-light conditions, leader shoots of tall trees naturally experience more severe water stress than those of short trees. We hypothesize that the height dependence of shoot allometry detected reflects an integrated response to height-associated water stress, which contributes to successful crown expansion and height gain.     

Linking water stress effects on carbon partitioning by introducing a xylem circuit into L-PEACH

Background and Aims

Many physiological processes such as photosynthesis, respiration and transpiration can be strongly influenced by the diurnal patterns of within-tree water potential. Despite numerous experiments showing the effect of water potential on fruit-tree development and growth, there are very few models combining carbohydrate allocation with water transport. The aim of this work was to include a xylem circuit into the functional–structural L-PEACH model.

Methods

The xylem modelling was based on an electrical circuit analogy and the Hagen–Poisseuille law for hydraulic conductance. Sub-models for leaf transpiration, soil water potential and the soil–plant interface were also incorporated to provide the driving force and pathway for water flow. The model was assessed by comparing model outputs to field measurements and published knowledge.

Key Results

The model was able to simulate both the water uptake over a season and the effect of different irrigation treatments on tree development, growth and fruit yield.

Conclusions

This work opens the way to a new field of modelling where complex interactions between water transport, carbohydrate allocation and physiological functions can be simulated at the organ level and describe functioning and behaviour at the tree scale.     

Stem hydraulic traits and leaf water-stress tolerance are co-ordinated with the leaf phenology of angiosperm trees in an Asian tropical dry karst forest

Background and Aims

The co-occurring of evergreen and deciduous angiosperm trees in Asian tropical dry forests on karst substrates suggests the existence of different water-use strategies among species. In this study it is hypothesized that the co-occurring evergreen and deciduous trees differ in stem hydraulic traits and leaf water relationships, and there will be correlated evolution in drought tolerance between leaves and stems.

Methods

A comparison was made of stem hydraulic conductivity, vulnerability curves, wood anatomy, leaf life span, leaf pressure–volume characteristics and photosynthetic capacity of six evergreen and six deciduous tree species co-occurring in a tropical dry karst forest in south-west China. The correlated evolution of leaf and stem traits was examined using both traditional and phylogenetic independent contrasts correlations.

Key Results

It was found that the deciduous trees had higher stem hydraulic efficiency, greater hydraulically weighted vessel diameter (D_h) and higher mass-based photosynthetic rate (A_m); while the evergreen species had greater xylem-cavitation resistance, lower leaf turgor-loss point water potential (π₀) and higher bulk modulus of elasticity. There were evolutionary correlations between leaf life span and stem hydraulic efficiency, A_m, and dry season π₀. Xylem-cavitation resistance was evolutionarily correlated with stem hydraulic efficiency, D_h, as well as dry season π₀. Both wood density and leaf density were closely correlated with leaf water-stress tolerance and A_m.

Conclusions

The results reveal the clear distinctions in stem hydraulic traits and leaf water-stress tolerance between the co-occurring evergreen and deciduous angiosperm trees in an Asian dry karst forest. A novel pattern was demonstrated linking leaf longevity with stem hydraulic efficiency and leaf water-stress tolerance. The results show the correlated evolution in drought tolerance between stems and leaves.Key words: Tropical dry forest, karst, leaf habit, hydraulic conductivity, cavitation resistance, leaf water-stress tolerance, wood density, leaf density, phylogenetic independent contrasts     

Plasticity of stomatal distribution pattern and stem tracheid dimensions in Podocarpus lambertii: an ecological study

Background and Aims

Leaf and wood plasticity are key elements in the survival of widely distributed plant species. Little is known, however, about variation in stomatal distribution in the leaf epidermis and its correlation with the dimensions of conducting cells in wood. This study aimed at testing the hypothesis that Podocarpus lambertii, a conifer tree, possesses a well-defined pattern of stomatal distribution, and that this pattern can vary together with the dimensions of stem tracheids as a possible strategy to survive in climatically different sites.

Methods

Leaves and wood were sampled from trees growing in a cold, wet site in south-eastern Brazil and in a warm, dry site in north-eastern Brazil. Stomata were thoroughly mapped in leaves from each study site to determine a spatial sampling strategy. Stomatal density, stomatal index and guard cell length were then sampled in three regions of the leaf: near the midrib, near the leaf margin and in between the two. This sampling strategy was used to test for a pattern and its possible variation between study sites. Wood and stomata data were analysed together via principal component analysis.

Key Results

The following distribution pattern was found in the south-eastern leaves: the stomatal index was up to 25 % higher in the central leaf region, between the midrib and the leaf margin, than in the adjacent regions. The inverse pattern was found in the north-eastern leaves, in which the stomatal index was 10 % higher near the midrib and the leaf margin. This change in pattern was accompanied by smaller tracheid lumen diameter and length.

Conclusions

Podocarpus lambertii individuals in sites with higher temperature and lower water availability jointly regulate stomatal distribution in leaves and tracheid dimensions in wood. The observed stomatal distribution pattern and variation appear to be closely related to the placement of conducting tissue in the mesophyll.     

Effects of light quality on leaf morphogenesis of a heterophyllous amphibious plant, Rotala hippuris

Background and Aims

For heterophyllous amphibious plants that experience fluctuating water levels, it is critical to control leaf development precisely in response to environmental cues that can serve as a quantitative index of water depth. Light quality can serve as such a cue because the ratio of red light relative to far-red light (R/FR) increases and blue-light intensity decreases with increasing water depth. Growth experiments were conducted to examine how R/FR and blue-light intensity alter leaf morphology of a heterophyllous amphibious plant, Rotala hippuris.

Methods

Using combinations of far red (730 nm), red (660 nm) and blue (470 nm) light-emitting diodes (LEDs), growth experiments were used to quantitatively evaluate the effects of the R/FR ratio and blue-light intensity on leaf morphology.

Key Results

Under the natural light regime in an outside growth garden, R. hippuris produced distinct leaves under submerged and aerial conditions. R/FR and blue-light intensity were found to markedly affect heterophyllous leaf formation. Higher and lower R/FR caused leaf characters more typical of submerged and aerial leaves, respectively, in both aerial and submerged conditions, in accordance with natural distribution of leaf types and light under water. High blue light caused a shift of trait values toward those of typical aerial leaves, and the response was most prominent under conditions of R/FR that were expected near the water surface.

Conclusions

R/FR and blue-light intensity provides quantitative cues for R. hippuris to detect water depth and determine the developmental fates of leaves, especially near the water surface. The utilization of these quantitative cues is expected to be important in habitats where plants experience water-level fluctuation.     

Genotypic variation in yellow autumn leaf colours explains aphid load in silver birch

? It has been suggested that autumn-migrating insects drive the evolution of autumn leaf colours. However, evidence of genetic variation in autumn leaf colours in natural tree populations and the link between the genetic variation and herbivore abundances has been lacking. ? Here, we measured the size of the whole aphid community and the development of green-yellow leaf colours in six replicate trees of 19 silver birch (Betula pendula) genotypes at the beginning, in the middle and at the end of autumn colouration. We also calculated the difference between green leaf and leaf litter nitrogen (N) and estimated the changes in phloem sap N loading. ? Autumn leaf colouration had significant genetic variation. During the last survey, genotypes that expressed the strongest leaf reflectance 2-4 wk earlier had an abundance of egg-laying Euceraphis betulae females. Surprisingly, the aphid community size during the first surveys explained N loss by the litter of different birch genotypes. ? Our results are the first evidence at the tree intrapopulation genotypic level that autumn-migrating pests have the potential to drive the evolution of autumn leaf colours. They also stress the importance of recognizing the role of late-season tree-insect interactions in the evolution of herbivory resistance.     

Fate and distribution of sulfur-35 in yellow poplar and red maple trees

Summary Two deciduous tree species (yellow poplar and red maple) on Walker Branch Watershed (WBW), near Oak Ridge, Tennessee, were radiolabeled with ³⁵S (87 day halflife) to study internal cycling, storage, and biogenic emission of sulfur (S). One tree of each species was girdled before radiolabeling to prevent phloem translocation to the roots, and the aboveground biomass was harvested prior to autumn leaf fall. Aboveground biomass, leaf fall, throughfall, and stemflow were sampled over a 13 to 24 week period. Sulfur-35 concentrations in tree leaves reached nearly asymptotic levels within 1 to 2 weeks after radiolabeling. Foliar leaching of ³⁵S and leaf fall represented relatively unimportant return pathways to the forest soil. The final distribution of ³⁵S in the nongirdled trees indicated little aboveground storage of S in biomass and appreciable (>60%) capacity to cycle S either to the belowground system by means of translocation or to the atmosphere by means of biogenic S emissions. Losses of volatile ³⁵S were estimated from the amount of isotope missing (33%) in final inventories of the girdled trees. Estimated ³⁵S emission rates from the girdled trees were 10^-6 to 10^-7 Ci cm^-2 leaf d^-1, and corresponded to an estimated gaseous S emission of approximately 0.1 to 1 g S cm^-2 leaf d^-1. Translocation to roots was a significant sink for ³⁵S in the red maple tree (40% of the injected amount). Research on forest biogeochemical S cycles should further explore biogenic S emissions from trees as a potential process of S flux from forest ecosystems.     

Developmental changes in spatial distribution of in vivo fluorescence and epidermal UV absorbance over Quercus petraea leaves

Background and Aims

Epidermal phenolic compounds (mainly flavonoids) constitute a vital screen that protects the leaf from damage by natural ultraviolet (UV) radiation. The effectiveness of epidermal UV-screening depends on leaf anatomy, the content of UV-screening compounds and their spatial uniformity over the leaf area. To investigate in vivo the spatial pattern of the epidermal UV-screen during leaf development, a fluorescence imaging method was developed to map the epidermal UV-absorbance at a microscopic scale. This study was done on oak (Quercus petraea) leaves that were used as a model of woody dicotyledonous leaves.

Methods

The leaf development of 2-year-old trees, grown outdoors, was monitored, at a macroscopic scale, by in vivo measurements of chlorophyll content per unit area and epidermal UV-absorbance using two optical leaf-clip meters. The distribution of pigments within leaves was assessed in vivo spectroscopically. The microscopic images of UV-induced fluorescence and UV-absorbance acquired in vivo during leaf development were interpreted from spectral characteristics of leaves.

Key Results

At a macroscopic scale, epidermal UV-absorbance was high on the upper leaf side during leaf development, while it increased on the lower leaf side during leaf expansion and reached the adaxial value at maturity. At a microscopic scale, in immature leaves, for both leaf sides, the spatial distribution of epidermal UV-absorbance was heterogeneous, with a pattern depending on the flavonoid content of vacuoles in developing epidermal cells. At maturity, epidermal UV-absorbance was uniform.

Conclusions

The spatial pattern of epidermal UV-screen over the area of oak leaves is related to leaf anatomy during development. In vivo spectroscopy and fluorescence imaging of the leaf surface showed the distribution of pigments within the leaf and hence can provide a tool to monitor optically the leaf development in nature.Key words: Blue-green fluorescence, chlorophyll fluorescence, epidermis, flavonoids, leaf development, microscopic imaging, polyphenols, Quercus petraea     

Red (anthocyanic) leaf margins do not correspond to increased phenolic content in New Zealand Veronica spp.

Background and Aims

Red or purple coloration of leaf margins is common in angiosperms, and is found in approx. 25 % of New Zealand Veronica species. However, the functional significance of margin coloration is unknown. We hypothesized that anthocyanins in leaf margins correspond with increased phenolic content in leaf margins and/or the leaf entire, signalling low palatability or leaf quality to edge-feeding insects.

Methods

Five species of Veronica with red leaf margins, and six species without, were examined in a common garden. Phenolic content in leaf margins and interior lamina regions of juvenile and fully expanded leaves was quantified using the Folin–Ciocalteu assay. Proportions of leaf margins eaten and average lengths of continuous bites were used as a proxy for palatability.

Key Results

Phenolic content was consistently higher in leaf margins compared with leaf interiors in all species; however, neither leaf margins nor more interior tissues differed significantly in phenolic content with respects to margin colour. Mean phenolic content was inversely correlated with the mean length of continuous bites, suggesting effective deterrence of grazing. However, there was no difference in herbivore consumption of red and green margins, and the plant species with the longest continuous grazing patterns were both red-margined.

Conclusions

Red margin coloration was not an accurate indicator of total phenolic content in leaf margins or interior lamina tissue in New Zealand Veronica. Red coloration was also ineffective in deterring herbivory on the leaf margin, though studies controlling for variations in leaf structure and biochemistry (e.g. intra-specific studies) are needed before more precise conclusions can be drawn. It is also recommended that future studies focus on the relationship between anthocyanin and specific defence compounds (rather than general phenolic pools), and evaluate possible alternative functions of red margins in leaves (e.g. antioxidants, osmotic adjustment).     

Deciduous and evergreen trees differ in juvenile biomass allometries because of differences in allocation to root storage

Background and Aims

Biomass partitioning for resource conservation might affect plant allometry, accounting for a substantial amount of unexplained variation in existing plant allometry models. One means of resource conservation is through direct allocation to storage in particular organs. In this study, storage allocation and biomass allometry of deciduous and evergreen tree species from seasonal environments were considered. It was expected that deciduous species would have greater allocation to storage in roots to support leaf regrowth in subsequent growing seasons, and consequently have lower scaling exponents for leaf to root and stem to root partitioning, than evergreen species. It was further expected that changes to root carbohydrate storage and biomass allometry under different soil nutrient supply conditions would be greater for deciduous species than for evergreen species.

Methods

Root carbohydrate storage and organ biomass allometries were compared for juveniles of 20 savanna tree species of different leaf habit (nine evergreen, 11 deciduous) grown in two nutrient treatments for periods of 5 and 20 weeks (total dry mass of individual plants ranged from 0·003 to 258·724 g).

Key Results

Deciduous species had greater root non-structural carbohydrate than evergreen species, and lower scaling exponents for leaf to root and stem to root partitioning than evergreen species. Across species, leaf to stem scaling was positively related, and stem to root scaling was negatively related to root carbohydrate concentration. Under lower nutrient supply, trees displayed increased partitioning to non-structural carbohydrate, and to roots and leaves over stems with increasing plant size, but this change did not differ between leaf habits.

Conclusions

Substantial unexplained variation in biomass allometry of woody species may be related to selection for resource conservation against environmental stresses, such as resource seasonality. Further differences in plant allometry could arise due to selection for different types of biomass allocation in response to different environmental stressors (e.g. fire vs. herbivory).     

Optimal photosynthetic use of light by tropical tree crowns achieved by adjustment of individual leaf angles and nitrogen content

Background and Aims

Theory for optimal allocation of foliar nitrogen (ONA) predicts that both nitrogen concentration and photosynthetic capacity will scale linearly with gradients of insolation within plant canopies. ONA is expected to allow plants to efficiently use both light and nitrogen. However, empirical data generally do not exhibit perfect ONA, and light-use optimization per se is little explored. The aim was to examine to what degree partitioning of nitrogen or light is optimized in the crowns of three tropical canopy tree species.

Methods

Instantaneous photosynthetic photon flux density (PPFD) incident on the adaxial surface of individual leaves was measured along vertical PPFD gradients in tree canopies at a frequency of 0·5 Hz over 9–17 d, and summed to obtain the average daily integral of PPFD for each leaf to characterize its insolation regime. Also measured were leaf N per area (N_area), leaf mass per area (LMA), the cosine of leaf inclination and the parameters of the photosynthetic light response curve [photosynthetic capacity (A_max), dark respiration (R_d), apparent quantum yield (ϕ) and curvature (θ)]. The instantaneous PPFD measurements and light response curves were used to estimate leaf daily photosynthesis (A_daily) for each leaf.

Key Results

Leaf N_area and A_max changed as a hyperbolic asymptotic function of the PPFD regime, not the linear relationship predicted by ONA. Despite this suboptimal nitrogen partitioning among leaves, A_daily did increase linearly with PPFD regime through co-ordinated adjustments in both leaf angle and physiology along canopy gradients in insolation, exhibiting a strong convergence among the three species.

Conclusions

The results suggest that canopy tree leaves in this tropical forest optimize photosynthetic use of PPFD rather than N per se. Tropical tree canopies then can be considered simple ‘big-leaves’ in which all constituent ‘small leaves’ use PPFD with the same photosynthetic efficiency.Key words: Optimal resource allocation, nitrogen, photosynthetic capacity, leaf mass per area, tropical trees, radiation use efficiency, scaling, leaf angle, canopy architecture, big leaf model