Anthocyanins act as a sugar-buffer and an alternative electron sink in response to starch depletion during leaf senescence: a case study on a typical anthocyanic tree species, Acer japonicum.

We hypothesized that anthocyanins act as a sugar-buffer and an alternative electron sink during leaf senescence to prevent sugar-mediated early senescence and photoinhibition. To elucidate the role of anthocyanin, we monitored seasonal changes in photosynthetic traits, sugar, starch and N contents, pigment composition, and gene expression profiles in leaves exposed to substantially different light conditions within a canopy of an adult fullmoon maple (Acer japonicum) tree. Enhancement of starch amylolysis accompanied by cessation of starch synthesis occurred in the same manner independent of light conditions. Leaf sugar contents increased, but reached upper limits in the late stage of leaf senescence, even though leaf anthocyanins further increased after complete depletion of starch. Sun-exposed leaves maintained higher energy consumption via electron flow than shade-grown leaves during leaf N resorption. Thus, anthocyanins accumulated in sun-exposed leaves might have a regulative role as a sugar-buffer, retarding leaf senescence, and an indirect photoprotective role as an alternative sink for electron consumption to compensate declines in other metabolic processes such as starch and protein synthesis. In this context, anthocyanins may be key substrates protecting both outer-canopy leaves (against photoinhibition) and inner-canopy leaves (via shading by outer-canopy leaves) from high light stress during N resorption.

Urbanization exacerbates climate sensitivity of eastern United States broadleaf trees.

Tree growth is a key mechanism driving carbon sequestration in forest ecosystems. Environmental conditions are important regulators of tree growth that can vary considerably between nearby urban and rural forests. For example, trees growing in cities often experience hotter and drier conditions than their rural counterparts while also being exposed to higher levels of light, pollution, and nutrient inputs. However, the extent to which these intrinsic differences in the growing conditions of trees in urban versus rural forests influence tree growth response to climate is not well known. In this study, we tested for differences in the climate sensitivity of tree growth between urban and rural forests along a latitudinal transect in the eastern United States that included Boston, Massachusetts, New York City, New York, and Baltimore, Maryland. Using dendrochronology analyses of tree cores from 55 white oak trees (Quercus alba), 55 red maple trees (Acer rubrum), and 41 red oak trees (Quercus rubra) we investigated the impacts of heat stress and water stress on the radial growth of individual trees. Across our three-city study, we found that tree growth was more closely correlated with climate stress in the cooler climate cities of Boston and New York than in Baltimore. Furthermore, heat stress was a significant hindrance to tree growth in higher latitudes while the impacts of water stress appeared to be more evenly distributed across latitudes. We also found that the growth of oak trees, but not red maple trees, in the urban sites of Boston and New York City was more adversely impacted by heat stress than their rural counterparts, but we did not see these urban-rural differences in Maryland. Trees provide a wide range of important ecosystem services and increasing tree canopy cover was typically an important component of urban sustainability strategies. In light of our findings that urbanization can influence how tree growth responds to a warming climate, we suggest that municipalities consider these interactions when developing their tree-planting palettes and when estimating the capacity of urban forests to contribute to broader sustainability goals in the future.

No evidence of age-related decline in propagated Acer pseudoplatanus and Fraxinus excelsior plants.

Although a substantial body of evidence suggests that large and old trees have reduced metabolic levels, the search for the causes behind this observation has proved elusive. The strong coupling between age and size, commonly encountered in the field, precludes the isolation of the potential causes. We used standard propagation techniques (grafting and air-layering) to decouple the effects of size from those of age in affecting leaf structure, biochemistry and physiology of two broadleaved trees, Acer pseudoplatanus (a diffuse-porous species) and Fraxinus excelsior (a ring-porous species). The first year after establishment of the propagated plants, some of the measurements suggested the presence of age-related declines in metabolism, while other measurements either did not show any difference or suggested variability across treatments not associated with either age or size. During the second year after establishment, only one of the measured properties (specific leaf area) continued to show some evidence of an age-mediated decline (although much reduced compared to the field), whereas, for some properties (particularly for F. excelsior), even the opposite trend of age-related increases was apparent. We concluded that (1) our plants suffered from grafting shock during year 1 and they gradually recovered during year 2; (2) the results over 2 years do not support the statement that age directly mediates ageing in either species but instead suggest that size directly mediates ageing processes; and (3) neither shoots nor roots of A. pseudoplatanus showed any evidence of senescence.

Distinct redox state regulation in the seedling performance of Norway maple and sycamore.

Norway maple and sycamore, two Acer genus species, have an important ecological value and different sensitivity to stressing factors being currently aggravated by climate change. Seedling growth is postulated to be the main barrier for successful plant establishment under the climate change scenarios. Therefore, the differences in redox regulation during the seedling performance of Norway maple and sycamore were investigated. Seeds of the two Acer species exhibited an identical high germination capacity, whereas seedling emergence was higher in sycamores. PCA analyses revealed that there is more diversification in the leaf characteristics than roots. Norway maple displayed a higher chlorophyll content index (CCI) with a similar leaf mass whereas sycamore seedlings exhibited a higher normalized difference vegetation index (NDVI), higher water content, higher root biomass and higher shoot height. Based on NDVI, sycamore seedlings appeared as very healthy plants, whereas Norway maple seedlings displayed a moderate healthy phenotype. Therefore, redox basis of seedling performance was investigated. The total pool of glutathione was four times higher in sycamore leaves than in Norway maple leaves and was reflected in highly reduced half-cell reduction potential of glutathione. Sycamore leaves contained more ascorbate because the content of its reduced form (AsA) was twice as high as in Norway maple. Therefore, the AsA/DHA ratio was balanced in sycamore leaves, reaching 1, and was halved in Norway maple leaves. Nicotinamide adenine dinucleotide phosphate content was twice as high in sycamore leaves than in Norway maples; however, its reduced form (NADPH) was predominant in Norway maple seedlings. Norway maple leaves exhibited the highest anabolic and catabolic redox charge. The higher reduction capacity and the activity of NADPH-dependent reductases in Norway maple leaves possibly resulted in higher CCI, whereas the larger root system contributed to higher NDVI in sycamore. The different methods of controlling redox parameters in Acer seedlings grown at controlled conditions provided here can be useful in understanding how tree species can cope with a changing environment in the future.

Anatomical acclimation of mature leaves to increased irradiance in sycamore maple (Acer pseudoplatanus L.).

Trees regenerating in the understory respond to increased availability of light caused by gap formation by undergoing a range of morphological and physiological adjustments. These adjustments include the production of thick, sun-type leaves containing thicker mesophyll and longer palisade cells than in shade-type leaves. We asked whether in the shade-regenerating tree Acer pseudoplatanus, the increase in leaf thickness and expansion of leaf tissues are possible also in leaves that are already fully formed, a response reported so far only for a handful of species. We acclimated potted seedlings to eight levels (from 1 to 100%) of solar irradiance and, in late summer, transferred a subset of them to full sunlight. Within 30 days, the pre-shaded leaves increased leaf mass per area and became thicker mostly due to the elongation of palisade cells, except for the most shaded individuals which suffered irreversible photo-oxidative damage. This anatomical acclimation was accompanied by a transient decline in photosynthetic efficiency of PSII (Fv/FM), the magnitude of which was related to the degree of pre-shading. The Fv/FM recovered substantially within the re-acclimation period. However, leaves of transferred plants were shed earlier in the fall, indicating that the acclimation was not fully effective. These results show that A. pseudoplatanus is one of the few known species in which mature leaves may re-acclimate anatomically to increased irradiance. This may be an important mechanism enhancing utilization of gaps created during the growing season.

The xylem of anisohydric Quercus alba L. is more vulnerable to embolism than isohydric codominants.

The coordination of plant leaf water potential (ΨL ) regulation and xylem vulnerability to embolism is fundamental for understanding the tradeoffs between carbon uptake and risk of hydraulic damage. There is a general consensus that trees with vulnerable xylem more conservatively regulate ΨL than plants with resistant xylem. We evaluated if this paradigm applied to three important eastern US temperate tree species, Quercus alba L., Acer saccharum Marsh. and Liriodendron tulipifera L., by synthesizing 1600 ΨL observations, 122 xylem embolism curves and xylem anatomical measurements across 10 forests spanning pronounced hydroclimatological gradients and ages. We found that, unexpectedly, the species with the most vulnerable xylem (Q. alba) regulated ΨL less strictly than the other species. This relationship was found across all sites, such that coordination among traits was largely unaffected by climate and stand age. Quercus species are perceived to be among the most drought tolerant temperate US forest species; however, our results suggest their relatively loose ΨL regulation in response to hydrologic stress occurs with a substantial hydraulic cost that may expose them to novel risks in a more drought-prone future.

Friendly neighbours: Hydraulic redistribution accounts for one quarter of water used by neighbouring drought stressed tree saplings.

Hydraulic redistribution (HR) can buffer drought events of tree individuals, however, its relevance for neighbouring trees remains unclear. Here, we quantified HR to neighbouring trees in single- and mixed-species combinations. We hypothesized that uptake of HR water positively correlates with root length, number of root tips and root xylem hydraulic conductivity and that neighbours in single-species combinations receive more HR water than in phylogenetic distant mixed-species combinations. In a split-root experiment, a sapling with its roots split between two pots redistributed deuterium labelled water from a moist to a dry pot with an additional tree each. We quantified HR water received by the sapling in the dry pot for six temperate tree species. After 7 days, one quarter of the water in roots (2.1 ± 0.4 ml), stems (0.8 ± 0.2 ml) and transpiration (1.0 ± 0.3 ml) of the drought stressed sapling originated from HR. The amount of HR water transpired by the receiving plant stayed constant throughout the experiment. While the uptake of HR water increased with root length, species identity did not affect HR as saplings of Picea abies ((L.) Karst) and Fagus sylvatica (L.) in single- and mixed-species combinations received the same amount of HR water.

NAD(P)-Driven Redox Status Contributes to Desiccation Tolerance in Acer seeds.

Desiccation tolerance is a developmental program enabling seed survival in a dry state and is common in seeds categorized as orthodox. We focused on NAD and its phosphorylated form (NADP) because their continual switching between reduced (NAD(P)H) and oxidized (NAD(P)+) forms is involved in the modulation of redox signaling and the determination of the reducing power and further antioxidant responses. Norway maple and sycamore seeds representing the orthodox and recalcitrant categories, respectively, were used as models in a comparison of responses to water loss. The process of desiccation up to 10% water content (WC) was monitored in Norway maple seeds, while dehydration up to 30% WC was monitored in desiccation-sensitive sycamore seeds. Norway maple and sycamore seeds, particularly their embryonic axes, exhibited a distinct redox status during dehydration and desiccation. High NADPH levels, NAD+ accumulation, low and stable NAD(P)H/NAD(P)+ ratios expressed as reducing power and high NADPH-dependent enzyme activity were reported in Norway maple seeds and were considered attributes of orthodox-type seeds. The contrasting results of sycamore seeds contributed to their low antioxidant capacity and high sensitivity to desiccation. NADPH deficiency, low NADPH-dependent enzyme activity and lack of NAD+ accumulation were primary features of sycamore seeds, with implications for their NAD(P)H/NAD(P)+ ratios and reducing power and with effects on many seed traits. Thus, we propose that the distinct levels of pyridine nucleotides and their redox status contribute to orthodox and recalcitrant phenotype differentiation in seeds by affecting cellular redox signaling, metabolism and the antioxidant system.

Involvement of the MetO/Msr System in Two Acer Species That Display Contrasting Characteristics during Germination.

The levels of methionine sulfoxide (MetO) and the abundances of methionine sulfoxide reductases (Msrs) were reported as important for the desiccation tolerance of Acer seeds. To determine whether the MetO/Msrs system is related to reactive oxygen species (ROS) and involved in the regulation of germination in orthodox and recalcitrant seeds, Norway maple and sycamore were investigated. Changes in water content, MetO content, the abundance of MsrB1 and MsrB2 in relation to ROS content and the activity of reductases depending on nicotinamide adenine dinucleotides were monitored. Acer seeds differed in germination speed-substantially higher in sycamore-hydration dynamics, levels of hydrogen peroxide, superoxide anion radicals (O2•-) and hydroxyl radicals (•OH), which exhibited peaks at different stages of germination. The MetO level dynamically changed, particularly in sycamore embryonic axes, where it was positively correlated with the levels of O2•- and the abundance of MsrB1 and negatively with the levels of •OH and the abundance of MsrB2. The MsrB2 abundance increased upon sycamore germination; in contrast, it markedly decreased in Norway maple. We propose that the ROS-MetO-Msr redox system, allowing balanced Met redox homeostasis, participates in the germination process in sycamore, which is characterized by a much higher speed compared to Norway maple.

Transcriptome analysis based on a combination of sequencing platforms provides insights into leaf pigmentation in Acer rubrum.

BACKGROUND: Red maple (Acer rubrum L.) is one of the most common and widespread trees with colorful leaves. We found a mutant with red, yellow, and green leaf phenotypes in different branches, which provided ideal materials with the same genetic relationship, and little interference from the environment, for the study of complex metabolic networks that underly variations in the coloration of leaves. We applied a combination of NGS and SMRT sequencing to various red maple tissues. RESULTS: A total of 125,448 unigenes were obtained, of which 46 and 69 were thought to be related to the synthesis of anthocyanins and carotenoids, respectively. In addition, 88 unigenes were presumed to be involved in the chlorophyll metabolic pathway. Based on a comprehensive analysis of the pigment gene expression network, the mechanisms of leaf color were investigated. The massive accumulation of Cy led to its higher content and proportion than other pigments, which caused the redness of leaves. Yellow coloration was the result of the complete decomposition of chlorophyll pigments, the unmasking of carotenoid pigments, and a slight accumulation of Cy. CONCLUSIONS: This study provides a systematic analysis of color variations in the red maple. Moreover, mass sequence data obtained by deep sequencing will provide references for the controlled breeding of red maple.

Insights from in vivo micro-CT analysis: testing the hydraulic vulnerability segmentation in Acer pseudoplatanus and Fagus sylvatica seedlings.

The seedling stage is the most susceptible one during a tree's life. Water relations may be crucial for seedlings due to their small roots, limited water buffers and the effects of drought on water transport. Despite obvious relevance, studies on seedling xylem hydraulics are scarce as respective methodical approaches are limited. Micro-CT scans of intact Acer pseudoplatanus and Fagus sylvatica seedlings dehydrated to different water potentials (Ψ) allowed the simultaneous observation of gas-filled versus water-filled conduits and the calculation of percentage loss of conductivity (PLC) in stems, roots and leaves (petioles or main veins). Additionally, anatomical analyses were performed and stem PLC measured with hydraulic techniques. In A. pseudoplatanus, petioles showed a higher Ψ at 50% PLC (Ψ50 -1.13MPa) than stems (-2.51 MPa) and roots (-1.78 MPa). The main leaf veins of F. sylvatica had similar Ψ50 values (-2.26 MPa) to stems (-2.74 MPa) and roots (-2.75 MPa). In both species, no difference between root and stems was observed. Hydraulic measurements on stems closely matched the micro-CT based PLC calculations. Micro-CT analyses indicated a species-specific hydraulic architecture. Vulnerability segmentation, enabling a disconnection of the hydraulic pathway upon drought, was observed in A. pseudoplatanus but not in the especially shade-tolerant F. sylvatica. Hydraulic patterns could partly be related to xylem anatomical traits.

Synchrony in fall leaf drop: chlorophyll degradation, color change, and abscission layer formation in three temperate deciduous tree species.

PREMISE OF THE STUDY: Deciduous tree species remove some nutrients from their leaves during fall leaf senescence through retranslocation. Retranslocation impacts the timeline of leaf fall, amount of active chlorophyll, and overall leaf nitrogen content as fall color change occurs. Our objective was to identify interspecific differences in the timing of abscission layer formation, leaf color change, and the level of chlorophyll degradation of young trees during fall senescence. METHODS: Leaf relative chlorophyll content for three tree species was measured during fall 2015 by a greenness meter. These measurements were calibrated for each species through spectrophotometric determination of leaf chlorophyll concentration. Abscission layer formation was tracked using light microscopy of sampled leaves. Excised leaves were photographed on a flat white surface to track species leaf color through time. KEY RESULTS: All three species had different chlorophyll declination rates throughout the fall season. The maple species started with less chlorophyll and began abscission layer formation earlier. The other two species had a similar starting chlorophyll level and onset timing of abscission layer formation. Visible leaf color change was not associated with a threshold in either chlorophyll degradation or abscission layer formation across species. CONCLUSIONS: Maple species degraded less chlorophyll on average, in the fall, than did the oak and beech species. The rate of chlorophyll degradation in coordination with abscission layer formation varied by species. Color change was not a good predictor of level of chlorophyll degradation in leaves across species.

Frost fatigue response to simulated frost drought using a centrifuge in Acer mono Maxim.

In the coldest part of winter, water uptake is blocked by the frozen soil and frozen stems known as 'frost drought' causing severe embolisms in woody plants. Frost drought in stems was simulated in a centrifuge by a synergy between freeze-thaw cycles and the different tensions induced by changing the rotation speed. Frost fatigue was defined as a reduction of embolism resistance after a freeze-thaw cycle and determined from 'vulnerability curves', which showed percent losses of conductivity vs tension (positive value) or xylem pressure (negative value). Different tensions combined with a controlled freeze-thaw cycle were induced to investigate the effects on frost resistance over the course of year. During the growing season, Acer mono Maxim. developed significant frost fatigue, and a significant positive correlation was found between frost fatigue response and exogenous tension. During the dormant season, A. mono showed very high embolism resistance to frost drought, even under a tension of 2 MPa. When the exogenous tension was increased to 3 MPa while the stem was frozen, significant frost fatigue occurred. Longer freezing times had more serious effects on frost fatigue in A. mono. A hypothesis was raised that at the same low temperature, the severer the drought (higher tension) when stems were frozen, the higher frost fatigue response would be induced.

Hydraulic safety margins and air-seeding thresholds in roots, trunks, branches and petioles of four northern hardwood trees.

During drought, xylem sap pressures can approach or exceed critical thresholds where gas embolisms form and propagate through the xylem network, leading to systemic hydraulic dysfunction. The vulnerability segmentation hypothesis (VSH) predicts that low-investment organs (e.g. leaf petioles) should be more vulnerable to embolism spread compared to high-investment, perennial organs (e.g. trunks, stems), as a means of mitigating embolism spread and excessive negative pressures in the perennial organs. We tested this hypothesis by measuring air-seeding thresholds using the single-vessel air-injection method and calculating hydraulic safety margins in four northern hardwood tree species of the northeastern United States, in both saplings and canopy height trees, and at five points along the soil-plant-atmosphere continuum. Acer rubrum was the most resistant to air-seeding and generally supported the VSH. However, Fagus grandifolia, Fraxinus americana and Quercus rubra showed little to no variation in air-seeding thresholds across organ types within each species. Leaf-petiole xylem operated at water potentials close to or exceeding their hydraulic safety margins in all species, whereas roots, trunks and stems of A. rubrum, F. grandifolia and Q. rubra operated within their safety margins, even during the third-driest summer in the last 100 yr.

New possible mechanisms of embolism formation when measuring vulnerability curves by air injection in a pressure sleeve.

Since 1988, researchers have exposed stems to positive pressures to displace water in vessels and measure the impact of applied pressure on hydraulic conductivity. The pressure-sleeve technique has been used in more than 60 publications to measure vulnerability curves (VCs), which are a measure of how water stress impacts the ability of plants to transport water because water stress induces embolism in vessels that blocks water flow. It is thought that the positive pressure in a sleeve required to induce 50% loss of conductivity (PLC), P50 , is the same magnitude as the tension that causes 50% PLC, T50 , where the tension can be induced by either bench-top dehydration or by a centrifuge technique. The unifying concept that P50  = T50 and that the entire VC is the same regardless of method is referred to as the air-seeding hypothesis. In the current study, we performed experiments to further test the air-seeding hypothesis in pressure sleeves and concluded that an "effervescence" mechanism caused embolism formation under positive pressure. This mechanism explains why VCs measured using positive pressure do not always match VCs obtained by other methods that induce water tension.

Seasonality of cavitation and frost fatigue in Acer mono Maxim.

Although cavitation is common in plants, it is unknown whether the cavitation resistance of xylem is seasonally constant or variable. We tested the changes in cavitation resistance of Acer mono before and after a controlled cavitation-refilling and freeze-thaw cycles for a whole year. Cavitation resistance was determined from 'vulnerability curves' showing the percent loss of conductivity versus xylem tension. Cavitation fatigue was defined as a reduction of cavitation resistance following a cavitation-refilling cycle, whereas frost fatigue was caused by a freeze-thaw cycle. A. mono developed seasonal changes in native embolisms; values were relatively high during winter but relatively low and constant throughout the growing season. Cavitation fatigue occurred and changed seasonally during the 12-month cycle; the greatest fatigue response occurred during summer and the weakest during winter, and the transitions occurred during spring and autumn. A. mono was highly resistant to frost damage during the relatively mild winter months; however, a quite different situation occurred during the growing season, as the seasonal trend of frost fatigue was strikingly similar to that of cavitation fatigue. Seasonality changes in cavitation resistance may be caused by seasonal changes in the mechanical properties of the pit membranes.

A network model links wood anatomy to xylem tissue hydraulic behaviour and vulnerability to cavitation.

Plant xylem response to drought is routinely represented by a vulnerability curve (VC). Despite the significance of VCs, the connection between anatomy and tissue-level hydraulic response to drought remains a subject of inquiry. We present a numerical model of water flow in flowering plant xylem that combines current knowledge on diffuse-porous anatomy and embolism spread to explore this connection. The model produces xylem networks and uses different parameterizations of intervessel connection vulnerability to embolism spread: the Young-Laplace equation and pit membrane stretching. Its purpose is upscaling processes occurring on the microscopic length scales, such as embolism propagation through pit membranes, to obtain tissue-scale hydraulics. The terminal branch VC of Acer glabrum was successfully reproduced relying only on real observations of xylem tissue anatomy. A sensitivity analysis shows that hydraulic performance and VC shape and location along the water tension axis are heavily dependent on anatomy. The main result is that the linkage between pit-scale and vessel-scale anatomical characters, along with xylem network topology, affects VCs significantly. This work underscores the importance of stepping up research related to the three-dimensional network structure of xylem tissues. The proposed model's versatility makes it an important tool to explore similar future questions.

Adenylate control contributes to thermal acclimation of sugar maple fine-root respiration in experimentally warmed soil.

We investigated the occurrence of and mechanisms responsible for acclimation of fine-root respiration of mature sugar maple (Acer saccharum) after 3+ years of experimental soil warming (+4 to 5 °C) in a factorial combination with soil moisture addition. Potential mechanisms for thermal respiratory acclimation included changes in enzymatic capacity, as indicated by root N concentration; substrate limitation, assessed by examining nonstructural carbohydrates and effects of exogenous sugar additions; and adenylate control, examined as responses of root respiration to a respiratory uncoupling agent. Partial acclimation of fine-root respiration occurred in response to soil warming, causing specific root respiration to increase to a much lesser degree (14% to 26%) than would be expected for a 4 to 5 °C temperature increase (approximately 55%). Acclimation was greatest when ambient soil temperature was warmer or soil moisture availability was low. We found no evidence that enzyme or substrate limitation caused acclimation but did find evidence supporting adenylate control. The uncoupling agent caused a 1.4 times greater stimulation of respiration in roots from warmed soil. Sugar maple fine-root respiration in warmed soil was at least partially constrained by adenylate use, helping constrain respiration to that needed to support work being performed by the roots.

Linking fine root morphology, hydraulic functioning and shade tolerance of trees.

Background and Aims: Understanding root traits and their trade-off with other plant processes is important for understanding plant functioning in natural ecosystems as well as agricultural systems. The aim of the present study was to determine the relationship between root morphology and the hydraulic characteristics of several orders of fine roots (<2 mm) for species differing in shade tolerance (low, moderate and high). Methods: The morphological, anatomical and hydraulic traits across five distal root orders were measured in species with different levels of shade tolerance and life history strategies. The species studied were Acer negundo, Acer rubrum, Acer saccharum, Betula alleghaniensis, Betula lenta, Quercus alba, Quercus rubra, Pinus strobus and Pinus virginiana. Key Results: Compared with shade-tolerant species, shade-intolerant species produced thinner absorptive roots with smaller xylem lumen diameters and underwent secondary development less frequently, suggesting that they had shorter life spans. Shade-tolerant species had greater root specific hydraulic conductance among these roots due to having larger diameter xylems, although these roots had a lower calculated critical tension for conduit collapse. In addition, shade-intolerant species exhibited greater variation in hydraulic conductance across different root growth rings in woody transport roots of the same root order as compared with shade-tolerant species. Conclusions: Plant growth strategies were extended to include root hydraulic properties. It was found that shade intolerance in trees was associated with conservative root hydraulics but greater plasticity in number of xylem conduits and hydraulic conductance. Root traits of shade-intolerant species were consistent with the ability to proliferate roots quickly for rapid water uptake needed to support rapid shoot growth, while minimizing risk in uncertain environments.

Convergence in leaf size versus twig leaf area scaling: do plants optimize leaf area partitioning?

BACKGROUND AND AIMS: Corner's rule states that thicker twigs bear larger leaves. The exact nature of this relationship and why it should occur has been the subject of numerous studies. It is obvious that thicker twigs should support greater total leaf area ([Formula: see text]) for hydraulical and mechanical reasons. But it is not obvious why mean leaf size ([Formula: see text]) should scale positively with [Formula: see text] We asked what this scaling relationship is within species and how variable it is across species. We then developed a model to explain why these relationships exist. METHODS: To minimize potential sources of variability, we compared twig properties from six co-occurring and functionally similar species: Acer grandidentatum, Amelanchier alnifolia, Betula occidentalis, Cornus sericea, Populus fremontii and Symphoricarpos oreophilus We modelled the economics of leaf display, weighing the benefit from light absorption against the cost of leaf tissue, to predict the optimal [Formula: see text] combinations under different canopy openings. KEY RESULTS: We observed a common [Formula: see text] by [Formula: see text] exponent of 0.6, meaning that [Formula: see text]and leaf number on twigs increased in a specific coordination. Common scaling exponents were not supported for relationships between any other measured twig properties. The model consistently predicted positive [Formula: see text] by [Formula: see text] scaling when twigs optimally filled canopy openings. The observed 0·6 exponent was predicted when self-shading decreased with larger canopy opening. CONCLUSIONS: Our results suggest Corner's rule may be better understood when recast as positive [Formula: see text] by [Formula: see text] scaling. Our model provides a tentative explanation of observed [Formula: see text] by [Formula: see text] scaling and suggests different scaling may exist in different environments.