Global convergence in the vulnerability of forests to drought.

Shifts in rainfall patterns and increasing temperatures associated with climate change are likely to cause widespread forest decline in regions where droughts are predicted to increase in duration and severity. One primary cause of productivity loss and plant mortality during drought is hydraulic failure. Drought stress creates trapped gas emboli in the water transport system, which reduces the ability of plants to supply water to leaves for photosynthetic gas exchange and can ultimately result in desiccation and mortality. At present we lack a clear picture of how thresholds to hydraulic failure vary across a broad range of species and environments, despite many individual experiments. Here we draw together published and unpublished data on the vulnerability of the transport system to drought-induced embolism for a large number of woody species, with a view to examining the likely consequences of climate change for forest biomes. We show that 70% of 226 forest species from 81 sites worldwide operate with narrow (<1 megapascal) hydraulic safety margins against injurious levels of drought stress and therefore potentially face long-term reductions in productivity and survival if temperature and aridity increase as predicted for many regions across the globe. Safety margins are largely independent of mean annual precipitation, showing that there is global convergence in the vulnerability of forests to drought, with all forest biomes equally vulnerable to hydraulic failure regardless of their current rainfall environment. These findings provide insight into why drought-induced forest decline is occurring not only in arid regions but also in wet forests not normally considered at drought risk.

Vulnerability to xylem embolism as a major correlate of the environmental distribution of rain forest species on a tropical island.

Increases in drought-induced tree mortality are being observed in tropical rain forests worldwide and are also likely to affect the geographical distribution of tropical vegetation. However, the mechanisms underlying the drought vulnerability and environmental distribution of tropical species have been little studied. We measured vulnerability to xylem embolism (P50 ) of 13 woody species endemic to New Caledonia and with different xylem conduit morphologies. We examined the relation between P50 , along with other leaf and xylem functional traits, and a range of habitat variables. Selected species had P50 values ranging between -4.03 and -2.00 MPa with most species falling in a narrow range of resistance to embolism above -2.7 MPa. Embolism vulnerability was significantly correlated with elevation, mean annual temperature and percentage of species occurrences located in rain forest habitats. Xylem conduit type did not explain variation in P50 . Commonly used functional traits such as wood density and leaf traits were not related to embolism vulnerability. Xylem embolism vulnerability stands out among other commonly used functional traits as a major driver of species environmental distribution. Drought-induced xylem embolism vulnerability behaves as a physiological trait closely associated with the habitat occupation of rain forest woody species.

Weak tradeoff between xylem safety and xylem-specific hydraulic efficiency across the world's woody plant species.

The evolution of lignified xylem allowed for the efficient transport of water under tension, but also exposed the vascular network to the risk of gas emboli and the spread of gas between xylem conduits, thus impeding sap transport to the leaves. A well-known hypothesis proposes that the safety of xylem (its ability to resist embolism formation and spread) should trade off against xylem efficiency (its capacity to transport water). We tested this safety-efficiency hypothesis in branch xylem across 335 angiosperm and 89 gymnosperm species. Safety was considered at three levels: the xylem water potentials where 12%, 50% and 88% of maximal conductivity are lost. Although correlations between safety and efficiency were weak (r(2)  < 0.086), no species had high efficiency and high safety, supporting the idea for a safety-efficiency tradeoff. However, many species had low efficiency and low safety. Species with low efficiency and low safety were weakly associated (r(2)  < 0.02 in most cases) with higher wood density, lower leaf- to sapwood-area and shorter stature. There appears to be no persuasive explanation for the considerable number of species with both low efficiency and low safety. These species represent a real challenge for understanding the evolution of xylem.

Stable isotopes in leaf water of terrestrial plants.

Leaf water contains naturally occurring stable isotopes of oxygen and hydrogen in abundances that vary spatially and temporally. When sufficiently understood, these can be harnessed for a wide range of applications. Here, we review the current state of knowledge of stable isotope enrichment of leaf water, and its relevance for isotopic signals incorporated into plant organic matter and atmospheric gases. Models describing evaporative enrichment of leaf water have become increasingly complex over time, reflecting enhanced spatial and temporal resolution. We recommend that practitioners choose a model with a level of complexity suited to their application, and provide guidance. At the same time, there exists some lingering uncertainty about the biophysical processes relevant to patterns of isotopic enrichment in leaf water. An important goal for future research is to link observed variations in isotopic composition to specific anatomical and physiological features of leaves that reflect differences in hydraulic design. New measurement techniques are developing rapidly, enabling determinations of both transpired and leaf water δ(18) O and δ(2) H to be made more easily and at higher temporal resolution than previously possible. We expect these technological advances to spur new developments in our understanding of patterns of stable isotope fractionation in leaf water.

Intercontinental long-distance dispersal of Canellaceae from the New to the Old World revealed by a nuclear single copy gene and chloroplast loci.

Canellales, a clade consisting of Winteraceae and Canellaceae, represent the smallest order of magnoliid angiosperms. The clade shows a broad distribution throughout the Southern Hemisphere, across a diverse range of dry to wet tropical forests. In contrast to their sister-group, Winteraceae, the phylogenetic relations and biogeography within Canellaceae remain poorly studied. Here we present the phylogenetic relationships of all currently recognized genera of Canellales with a special focus on the Old World Canellaceae using a combined dataset consisting of the chloroplast trnK-matK-trnK-psbA and the nuclear single copy gene mag1 (Maigo 1). Within Canellaceae we found high statistical support for the monophyly of Warburgia and Cinnamosma. However, we also found relationships that differ from previous studies. Cinnamodendron splitted into two clades, a South American clade and a second clade confined to the Antilles and adjacent areas. Cinnamodendron from the Antilles, as well as Capsicodendron, South American Cinnamodendron and Pleodendron were not monophyletic. Consequently, Capsicodendron should be included in the South American Cinnamodendron clade and the genus Pleodendron merged with the Cinnamodendron clade from the Antilles. We also found that Warburgia (restricted to mainland eastern Africa) together with the South American Cinnamodendron and Capsicodendron are sister to the Malagasy genus Cinnamosma. In addition to the unexpected geographical relationships, both biogeographic and molecular clock analyses suggest vicariance, extinction, and at least one intercontinental long-distance-dispersal event. Our dating result contrasts previous work on Winteraceae. Diversification of Winteraceae took place in the Paleocene, predating the Canellaceae diversification by 13 MA in the Eocene. The phylogenetic relationships for Canellaceae supported here offer a solid framework for a future taxonomic revision of the Canellaceae.

Assessing the impact of phylogenetic incongruence on taxonomy, floral evolution, biogeographical history, and phylogenetic diversity.

PREMISE OF THE STUDY: Phylogenetic incongruence between "gene trees" and "species trees" has been widely acknowledged in phylogenetic research. Conflicts may emerge from several processes including paralogy, hybridization, and incomplete lineage sorting. Although phylogenetic incongruence appears common, its impact on many phylogeny-based analyses remains poorly understood. METHODS: We examined the occurrence of phylogenetic conflict between nuclear (ribosome ITS) and plastid (rbcL, trnL-F, rpl20-rps12, and rps16 intron) loci in the ancient angiosperm family Chloranthaceae. Then we investigated how phylogenetic conflict bears on taxonomic classification within the family as well as on inferences on biogeographical history, floral evolution, and measures of phylogenetic diversity (PD). KEY RESULTS: We found evidence for significant phylogenetic incongruence between plastid and nuclear data in the genus Hedyosmum. Within Hedyosmum, our results did not support previous subgeneric classification of the genus. Division of sections within subgenus Tafalla was supported by the ITS data but not by the plastid data set. As a consequence, we showed that inferring the evolution of key floral characters and geographical history within Hedyosmum depends on the phylogenetic data used. Both data sets yielded similar PD measures across genera, but we found contrasting PD measures in Hedyosmum, even after correcting for rate heterogeneity. CONCLUSIONS: Our study demonstrated that phylogenetic conflict not only affects the inference of organismal relationships but also impacts our understanding of biogeographical history, morphological evolution, and phylogenetic diversity.

Paleo-Antarctic rainforest into the modern Old World tropics: the rich past and threatened future of the "southern wet forest survivors".

UNLABELLED: • PREMISE OF STUDY: Have Gondwanan rainforest floral associations survived? Where do they occur today? Have they survived continuously in particular locations? How significant is their living floristic signal? We revisit these classic questions in light of significant recent increases in relevant paleobotanical data.• METHODS: We traced the extinction and persistence of lineages and associations through the past across four now separated regions-Australia, New Zealand, Patagonia, and Antarctica-using fossil occurrence data from 63 well-dated Gondwanan rainforest sites and 396 constituent taxa. Fossil sites were allocated to four age groups: Cretaceous, Paleocene-Eocene, Neogene plus Oligocene, and Pleistocene. We compared the modern and ancient distributions of lineages represented in the fossil record to see if dissimilarity increased with time. We quantified similarity-dissimilarity of composition and taxonomic structure among fossil assemblages, and between fossil and modern assemblages.• KEY RESULTS: Strong similarities between ancient Patagonia and Australia confirmed shared Gondwanan rainforest history, but more of the lineages persisted in Australia. Samples of ancient Australia grouped with the extant floras of Australia, New Guinea, New Caledonia, Fiji, and Mt. Kinabalu. Decreasing similarity through time among the regional floras of Antarctica, Patagonia, New Zealand, and southern Australia reflects multiple extinction events.• CONCLUSIONS: Gondwanan rainforest lineages contribute significantly to modern rainforest community assembly and often co-occur in widely separated assemblages far from their early fossil records. Understanding how and where lineages from ancient Gondwanan assemblages co-occur today has implications for the conservation of global rainforest vegetation, including in the Old World tropics.

Fossil evidence for Cretaceous escalation in angiosperm leaf vein evolution.

The flowering plants that dominate modern vegetation possess leaf gas exchange potentials that far exceed those of all other living or extinct plants. The great divide in maximal ability to exchange CO(2) for water between leaves of nonangiosperms and angiosperms forms the mechanistic foundation for speculation about how angiosperms drove sweeping ecological and biogeochemical change during the Cretaceous. However, there is no empirical evidence that angiosperms evolved highly photosynthetically active leaves during the Cretaceous. Using vein density (D(V)) measurements of fossil angiosperm leaves, we show that the leaf hydraulic capacities of angiosperms escalated several-fold during the Cretaceous. During the first 30 million years of angiosperm leaf evolution, angiosperm leaves exhibited uniformly low vein D(V) that overlapped the D(V) range of dominant Early Cretaceous ferns and gymnosperms. Fossil angiosperm vein densities reveal a subsequent biphasic increase in D(V). During the first mid-Cretaceous surge, angiosperm D(V) first surpassed the upper bound of D(V) limits for nonangiosperms. However, the upper limits of D(V) typical of modern megathermal rainforest trees first appear during a second wave of increased D(V) during the Cretaceous-Tertiary transition. Thus, our findings provide fossil evidence for the hypothesis that significant ecosystem change brought about by angiosperms lagged behind the Early Cretaceous taxonomic diversification of angiosperms.

Hydraulic tuning of vein cell microstructure in the evolution of angiosperm venation networks.

High vein density (D(V)) evolution in angiosperms represented a key functional transition. Yet, a mechanistic account on how this hydraulic transformation evolved remains lacking. We demonstrate that a consequence of producing high D(V is that veins must become very small to fit inside the leaf, and that angiosperms are the only clade that evolved the specific type of vessel required to yield sufficiently conductive miniature leaf veins. From 111 species spanning key divergences in vascular plant evolution, we show, using analyses of vein conduit evolution in relation to vein packing, that a key xylem innovation associated with high D(V) evolution is a strong reduction in vein thickness and simplification of the perforation plates of primary xylem vessels. Simple perforation plates in the leaf xylem occurred only in derived angiosperm clades exhibiting high D(V) (> 12 mm mm(-2)). Perforation plates in the vessels of other species, including extant basal angiosperms, consisted of resistive scalariform types that were associated with thicker veins and much lower D(V). We conclude that a reduction in within-vein conduit resistance allowed vein size to decrease. We suggest that this adaptation may have been a critical evolutionary step that enabled dramatic D(V) elaboration in angiosperms.

Uncorrelated evolution of leaf and petal venation patterns across the angiosperm phylogeny.

Early angiosperm evolution, beginning approximately 140 million years ago, saw many innovations that enabled flowering plants to alter ecosystems globally. These included the development of novel, flower-based pollinator attraction mechanisms and the development of increased water transport capacity in stems and leaves. Vein length per area (VLA) of leaves increased nearly threefold in the first 30-40 million years of angiosperm evolution, increasing the capacity for transpiration and photosynthesis. In contrast to leaves, high water transport capacities in flowers may not be an advantage because flowers do not typically contribute to plant carbon gain. Although flowers of extant basal angiosperms are hydrated by the xylem, flowers of more recently derived lineages may be hydrated predominantly by the phloem. In the present study, we measured leaf and flower VLA for a phylogenetically diverse sample of 132 species from 52 angiosperm families to ask (i) whether flowers have lower VLA than leaves, (ii) whether flowers of basal angiosperm lineages have higher VLA than more recently derived lineages because of differences between xylem and phloem hydration, and (iii) whether flower and leaf VLA evolved independently. It was found that floral structures had lower VLA than leaves, but basal angiosperm flowers did not have higher VLA than more derived lineages. Furthermore, the independent evolution of leaf and petal VLA suggested that these organs may be developmentally modular. Unlike leaves, which have experienced strong selection for increased water transport capacity, flowers may have been shielded from such selective pressures by different developmental processes controlling VLA throughout the plant bauplan.

The evolution of angiosperm lianescence without vessels--climbing mode and wood structure-function in Tasmannia cordata (Winteraceae).

• The lack of extant lianescent vessel-less seed plants supports a hypothesis that liana evolution requires large-diameter xylem conduits. Here, we demonstrate an unusual example of a lianoid vessel-less angiosperm, Tasmannia cordata (Winteraceae), from New Guinea. • Wood mechanical, hydraulic and structural measurements were used to determine how T. cordata climbs and to test for ecophysiological shifts related to liana evolution vs 13 free-standing congeners. • The tracheid-based wood of T. cordata furnished low hydraulic capacity compared with that of vessel-bearing lianas. In comparison with most nonclimbing relatives, T. cordata possessed lower photosynthetic rates and leaf and stem hydraulic capacities. However, T. cordata exhibited a two- to five-fold greater wood elastic modulus than its relatives. • Tasmannia cordata provides an unusual example of angiosperm liana evolution uncoupled from xylem conduit gigantism, as well as high plasticity and cell type diversity in vascular development. Because T. cordata lacks vessels, our results suggest that a key limitation for a vessel-less liana is that strong and low hydraulically conductive wood is required to meet the mechanical demands of lianescence.

Leaf hydraulic evolution led a surge in leaf photosynthetic capacity during early angiosperm diversification.

Angiosperm evolution transformed global ecology, and much of this impact derives from the unrivalled vegetative productivity of dominant angiosperm clades. However, the origins of high photosynthetic capacity in angiosperms remain unknown. In this study, we describe the steep trajectory of leaf vein density (D(v)) evolution in angiosperms, and predict that this leaf plumbing innovation enabled a major shift in the capacity of leaves to assimilate CO(2). Reconstructing leaf vein evolution from an examination of 504 angiosperm species we found a rapid three- to fourfold increase in D(v) occurred during the early evolution of angiosperms. We demonstrate how this major shift in leaf vein architecture potentially allowed the maximum photosynthetic capacity in angiosperms to rise above competing groups 140-100 Ma. Our data suggest that early terrestrial angiosperms produced leaves with low photosynthetic rates, but that subsequent angiosperm success is linked to a surge in photosynthetic capacity during their early diversification.

Association between winter anthocyanin production and drought stress in angiosperm evergreen species.

Leaves of many evergreen angiosperm species turn red under high light during winter due to the production of anthocyanin pigments, while leaves of other species remain green. There is currently no explanation for why some evergreen species exhibit winter reddening while others do not. Conditions associated with low leaf water potentials (Psi) have been shown to induce reddening in many plant species. Because evergreen species differ in susceptibility to water stress during winter, it is hypothesized that species which undergo winter colour change correspond with those that experience/tolerate the most severe daily declines in leaf Psi during winter. Six angiosperm evergreen species which synthesize anthocyanin in leaves under high light during winter and five species which do not were studied. Field Psi, pressure/volume curves, and gas exchange measurements were derived in summer (before leaf colour change had occurred) and winter. Consistent with the hypothesis, red-leafed species as a group had significantly lower midday Psi in winter than green-leafed species, but not during the summer when all the leaves were green. However, some red-leafed species showed midday declines similar to those of green-leafed species, suggesting that low Psi alone may not induce reddening. Pressure-volume curves also provided some evidence of acclimation to more negative water potentials by red-leafed species during winter (e.g. greater osmotic adjustment and cell wall hardening on average). However, much overlap in these physiological parameters was observed as well between red and green-leafed species, and some of the least drought-acclimated species were red-leafed. No difference was observed in transpiration (E) during winter between red and green-leaved species. When data were combined, only three of the six red-leafed species examined appeared physiologically acclimated to prolonged drought stress, compared to one of the five green-leafed species. This suggests that drought stress alone is not sufficient to explain winter reddening in evergreen angiosperms.

Angiosperm leaf vein evolution was physiologically and environmentally transformative.

The veins that irrigate leaves during photosynthesis are demonstrated to be strikingly more abundant in flowering plants than in any other vascular plant lineage. Angiosperm vein densities average 8 mm of vein per mm(2) of leaf area and can reach 25 mm mm(-2), whereas such high densities are absent from all other plants, living or extinct. Leaves of non-angiosperms have consistently averaged close to 2 mm mm(-2) throughout 380 million years of evolution despite a complex history that has involved four or more independent origins of laminate leaves with many veins and dramatic changes in climate and atmospheric composition. We further demonstrate that the high leaf vein densities unique to the angiosperms enable unparalleled transpiration rates, extending previous work indicating a strong correlation between vein density and assimilation rates. Because vein density is directly measurable in fossils, these correlations provide new access to the physiology of extinct plants and how they may have impacted their environments. First, the high assimilation rates currently confined to the angiosperms among living plants are likely to have been unique throughout evolutionary history. Second, the transpiration-driven recycling of water that is important for bolstering precipitation in modern tropical rainforests might have been significantly less in a world before the angiosperms.

Evolution of stomatal responsiveness to CO(2) and optimization of water-use efficiency among land plants.

The stomata of angiosperms respond to changes in ambient atmospheric concentrations of CO(2) (C(a)) in ways that appear to optimize water-use efficiency. It is unknown where in the history of land plants this important stomatal control mechanism evolved. Here, we test the hypothesis that major clades of plants have distinct stomatal sensitivities to C(a) reflecting a relatively recent evolution of water-use optimization in derived angiosperms. Responses of stomatal conductance (g(s)) to step changes between elevated, ambient and low C(a) (600, 380 and 100 micromol mol(-1), respectively) were compared in a phylogenetically and ecologically diverse range of higher angiosperms, conifers, ferns and lycopods. All species responded to low C(a) by increasing g(s) but only angiosperm stomata demonstrated a significant closing response when C(a) was elevated to 600 micromol mol(-1). As a result, angiosperms showed significantly greater increases in water-use efficiency under elevated C(a) than the other lineages. The data suggest that the angiosperms have mechanisms for detecting and responding to increases in C(a) that are absent from earlier diverging lineages, and these mechanisms impart a greater capacity to optimize water-use efficiency.

Evolutionary significance of a flat-leaved Pinus in Vietnamese rainforest.

Pines are generally absent from tropical rainforests. An important exception, Pinus krempfii, is a unique tree that bears flattened needles and competes with evergreen angiosperm trees in southern Vietnam. Here, the photosynthetic and hydraulic physiology of P. krempfii leaves were examined to determine whether this species departs from the widespread pattern of high-light-demanding photosynthetic physiology displayed in needle-leaved Pinus species. Maximum photosynthesis and light saturation of photosynthesis, as well as stem and leaf hydraulic efficiencies, were all very low in P. krempfii compared with other Pinus species. These characteristics were consistent with our observations of P. krempfii seedling regeneration under the forest canopy. By possessing shade tolerance coupled with the production of flattened leaves, P. krempfii has converged morphologically and physiologically with many genera of the southern hemisphere conifer family Podocarpaceae. This convergence extends to a key feature of leaf anatomy, the production of tubular sclereids in the leaf for radial transport of water from the vein to the margin. These observations suggest that few adaptive possibilities are open to conifers when moving into tropical rainforest, meaning that Pinus is forced into direct competition with southern hemisphere conifers for a narrow niche in the equatorial zone.

The Emergence of Earliest Angiosperms may be Earlier than Fossil Evidence Indicates.

Gaps between molecular ages and fossils undermine the validity of time-calibrated molecular phylogenies. An example of the time gap surrounds the age of angiosperm's origin. We calculate molecular ages of the earliest flowering plant lineages using 22 fossil calibrations (101 genera, 40 families). Our results reveal the origin of angiosperms at the late Permian, ~275 million years ago. Different prior probability curves of molecular age calculations on dense calibration point distributions had little effect on overall age estimates compared to the effects of altered calibration points. The same is true for reasonable root age constraints. We conclude that our age estimates based on multiple datasets, priors, and calibration points are robust and the true ages are likely between our extremes. Our results, when integrated with the ecophysiological evolution of early angiosperms, imply that the ecology of the earliest angiosperms is critical to understand the pre-Cretaceous evolution of flowering plants.

Hardly a relict: freezing and the evolution of vesselless wood in Winteraceae.

The Winteraceae are traditionally regarded as the least-specialized descendents of the first flowering plants, based largely on their lack of xylem vessels. Since vessels have been viewed as a key innovation for angiosperm diversification, Winteraceae have been portrayed as declining relicts, limited to wet forest habitats where their tracheid-based wood does not impose a significant hydraulic constraints. In contrast, phylogenetic analyses place Winteraceae among angiosperm clades with vessels, indicating that their vesselless wood is derived rather than primitive, whereas extension of the Winteraceae fossil record into the Early Cretaceous suggests a more complex ecological history than has been deduced from their current distribution. However, the selective regime and ecological events underlying the possible loss of vessels in Winteraceae have remained enigmatic. Here we examine the hypothesis that vessels were lost as an adaptation to freezing-prone environments in Winteraceae by measuring the responses of xylem water transport to freezing for a diverse group of Winteraceae taxa as compared to Canella winterana (Canellaceae, a close relative with vessels) and sympatric conifer taxa. We found that mean percent loss of xylem water transport capacity following freeze-thaw varied from 0% to 6% for Winteraceae species from freezing-prone temperate climates and approximately 20% in those taxa from tropical (nonfreezing) climates. Similarly, conifers exhibit almost no decrease in xylem hydraulic conductivity following freezing. In contrast, water transport in Canella stems is nearly 85% blocked after freeze-thaw. Although vessel-bearing wood of Canella possesses considerably greaterhydraulic capacity than Winteraceae, nearly 20% of xylem hydraulic conductance remains, a value that is comparable to the hydraulic capacity of vesselless Winteraceae xylem, if the proportion of hydraulic flow through vessels (modeled as ideal capillaries) is removed. Thus, the evolutionary removal of vessels may not necessarily require a deleterious shift to an ineffective vascular system. By integrating Winteraceae's phylogenetic relationships and fossil history with physiological and ecological observations, we suggest that, as ancestors of modern Winteraceae passed through temperate conditions present in Southern Gondwana during the Early Cretaceous, they were exposed to selective pressures against vessel-possession and returned to a vascular system relying on tracheids. These results suggest that the vesselless condition is advantageous in freezing-prone areas, which is supported by the strong bias in the ecological abundance of Winteraceae to wet temperate and tropical alpine habitats, rather than a retained feature from the first vesselless angiosperms. We believe that vesselless wood plays an important role in the ecological abundance of Winteraceae in Southern Hemisphere temperate environments by enabling the retention of leaves and photosynthesis in the face of frequent freeze-thaw events.

Stem water transport and freeze-thaw xylem embolism in conifers and angiosperms in a Tasmanian treeline heath.

The effect of freezing on stem xylem hydraulic conductivity and leaf chlorophyll a fluorescence was measured in 12 tree and shrub species from a treeline heath in Tasmania, Australia. Reduction in stem hydraulic conductivity after a single freeze-thaw cycle was minimal in conifers and the vessel-less angiosperm species Tasmannia lanceolata (Winteraceae), whereas mean loss of conductivity in vessel-forming angiosperms fell in the range 17-83%. A positive linear relationship was observed between percentage loss of hydraulic conductivity by freeze-thaw and the average conduit diameter across all 12 species. This supports the hypothesis that large-diameter vascular conduits have a greater likelihood of freeze-thaw cavitation because larger bubbles are produced, which are more likely to expand under tension. Leaf frost tolerances, as measured by a 50% loss of maximum PSII quantum yield, varied from -6 to -13°C, indicating that these species were more frost-sensitive than plants from northern hemisphere temperate forest and treeline communities. There was no evidence of a relationship between frost tolerance of leaves and the resilience of stem water transport to freezing, suggesting that low temperature survival and the resistance of stem water transport to freezing are independently evolving traits. The results of this study bear on the ecological importance of stem freezing in the southern hemisphere treeline zones.

A potential role for xylem-phloem interactions in the hydraulic architecture of trees: effects of phloem girdling on xylem hydraulic conductance.

We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.