Temperate Woody Species Research Articles

Leaf water relations and structural traits of four temperate woody species occurring in serpentine and non‐serpentine soil

AbstractLow water‐holding properties impose dry conditions on plants on serpentine soil. To test the hypothesis that leaf water relations are key plant characteristics to grow in serpentine soil, we compared these traits for four tree species (Quercus serrata, Clethra barbinervis, Magnolia obovata and Pieris japonica) growing in serpentine soil and brown forest (BF) soil. Despite a much lower soil moisture content, trees in the serpentine soil (S) plot showed similar predawn leaf water potential to trees in the BF plot in all species. Trees in the S plot showed higher drought tolerance or drought avoidance than trees in the BF plot. In the S plot, enhanced drought tolerance through osmoregulation was observed in Q. serrata and P. japonica, whereas decreased leaf capacitance was observed in C. barbinervis and Q. serrata. Decreased stomatal conductance, a typical way to avoid drought stress, was observed for serpentine C. barbinervis and M. obovata and midday leaf water potential was comparable between S and BF plots for C. barbinervis. Lamina dry mass per area was higher for C. barbinervis but lower for Q. serrata in the S plot. Wood density was higher in serpentine Q. serrata and M. obovata. On the other hand, all studied species showed lower leaf nitrogen concentration in the S plot. Our results suggest that (a) despite low water availability, the tress in the serpentine plot did not suffer drought stress, and (b) leaf water relations may be one of the traits that enabled trees to inhabit serpentine soil.

Frost Survival Mechanism of Vegetative Buds in Temperate Trees: Deep Supercooling and Extraorgan Freezing vs. Ice Tolerance.

In temperate climates, overwintering buds of trees are often less cold hardy than adjoining stem tissues or evergreen leaves. However, data are scarce regarding the freezing resistance (FR) of buds and the underlying functional frost survival mechanism that in case of supercooling can restrict the geographic distribution. Twigs of 37 temperate woody species were sampled in midwinter 2016 in the Austrian Inn valley. After assessment of FR, infrared-video-thermography and cryo-microscopy were used to study the freezing pattern in and around overwintering vegetative buds. Only in four species, after controlled ice nucleation in the stem (−1.6 ± 0.9°C) ice was observed to immediately invade the bud. These buds tolerated extracellular ice and were the most freezing resistant (−61.8°C mean LT50). In all other species (33), the buds remained supercooled and free of ice, despite a frozen stem. A structural ice barrier prevents ice penetration. Extraorgan ice masses grew in the stem and scales but in 50% of the species between premature supercooled leaves. Two types of supercooled buds were observed: in temporary supercooling buds (14 species) ice spontaneously nucleated at −20.5 ± 4,6°C. This freezing process appeared to be intracellular as it matched the bud killing temperature (−22.8°C mean LT50). This response rendered temporarily supercooled buds as least cold hardy. In 19 species, the buds remained persistently supercooled down to below the killing temperature without indication for the cause of damage. Although having a moderate midwinter FR of −31.6°C (LT50), some species within this group attained a FR similar to ice tolerant buds. The present study represents the first comprehensive overview of frost survival mechanisms of vegetative buds of temperate trees. Except for four species that were ice tolerant, the majority of buds survive in a supercooled state, remaining free of ice. In 50% of species, extraorgan ice masses harmlessly grew between premature supercooled leaves. Despite exposure to the same environmental demand, midwinter FR of buds varied intra-specifically between −17.0 and −90.0°C. Particularly, species, whose buds are killed after temporary supercooling, have a lower maximum FR, which limits their geographic distribution.

Daily Maximum Temperatures Induce Lagged Effects on Leaf Unfolding in Temperate Woody Species Across Large Elevational Gradients.

The timing of leaf unfolding in temperate woody species is predominantly controlled by the seasonal course of temperature in late winter and early spring. However, quantifying lagged temperature effects on spring phenology is still challenging. Here, we aimed at investigating lagged and potentially non-linear effects of daily maximum temperatures on the probability of leaf unfolding in temperate woody species growing across large elevational gradients. We analyzed 5280 observations of leaf-out time of four tree species (European beech, horse chestnut, European larch, Norway spruce) and one shrub species (common hazel) that were recorded by volunteers over 40 years at 42 locations in Switzerland. We used a case-crossover sampling design to match leaf-out dates with control dates (i.e., dates before or after leaf-out), and analyzed these data with conditional logistic regression accounting for lagged temperature effects over 60 days. Multivariate meta-analyses were used to synthesize lagged temperature and elevational effects on leaf unfolding across multiple phenological stations. Temperature effects on the probability of leaf unfolding were largest at relatively short lags (i.e., within ca. 10 days) and decreased with increasing lags. Short- to mid-term effects (i.e., within ca. 10 to 20 days) were larger for late-leafing species known to be photoperiod-sensitive (beech, Norway spruce). Temperature effects increased for the broadleaved species (horse chestnut, hazel, beech) with decreasing elevation, particularly within ca. 10 to 40 days, i.e., leaf unfolding occurs more rapidly at low elevations for a given daily maximum temperature. Our novel findings provide evidence of cumulative and long-term temperature effects on leaf unfolding, whereby the efficiency of relatively high temperatures to trigger leaf-out becomes higher shortly before bud burst. These lagged associations between temperature and leaf unfolding improve our understanding of phenological responses across temperate woody species with differing ecological requirements that occur along elevational gradients.

Ongoing seasonally uneven climate warming leads to earlier autumn growth cessation in deciduous trees.

Ongoing global warming is causing phenological shifts that affect photosynthesis and growth rates in temperate woody species. However, the effects of seasonally uneven climate warming-as is occurring in much of Europe, where the winter/spring months are warming twice as fast than the summer/autumn months-on autumn growth cessation (completion of overwintering buds) and leaf senescence, and possible carry-over effects between phenophases, remain under-investigated. We conducted experiments in which we exposed saplings of canopy and understory species to 4°C warming in winter/spring, summer/autumn, or all year to disentangle how the timing of bud break, bud set completion, and leaf senescence is affected by seasonally uneven warming. All-year warming led to significantly delayed leaf senescence, but advanced bud set completion; summer/autumn warming only delayed leaf senescence; and winter/spring warming advanced both bud set and senescence. The non-parallel effects of warming on bud completion and leaf senescence show that leaf senescence alone is an inadequate proxy for autumn growth cessation in trees and counterintuitively suggest that continued uneven seasonal warming will advance cessation of primary growth in autumn, even when leaf senescence is delayed. Phenological responses to warming treatments (earlier spring onset, later autumn senescence) were more than twice as high in understory species than in canopy species, which can partly be explained by the absence of carry-over effects among phenophases in the former group. This underscores the need to consider differences among plant functional types when forecasting the future behaviour of ecosystems.

Extreme winter warm event causes exceptionally early bud break for many woody species

AbstractIn many ecosystems, climate is changing faster during winter compared to other seasons. However, we lack basic information about the responses of many species to winter climate change, including extreme warm events. Extreme warm events may have particularly strong influences at the end of winter, when some species begin to break dormancy while the risk of freezing remains high. Here, we monitored bud burst of 101 temperate woody species following an extreme warm event during winter to investigate species responses to this anomalous event and determine whether functional traits predicted species responses. Following six consecutive days of extreme warm temperatures in winter, nearly half the surveyed tree and shrub species had an advanced stage of bud phenology. Responding species were most likely to be shade‐intolerant, phylogenetically related, and have weaker dormancy requirements. Based on established species response thresholds to spring temperatures in the region, species were expected to be unresponsive to warm temperatures this early in the year, yet many species broke dormancy. Species responses to this extreme winter warm event highlighted how climate change can alter well‐established species–climate associations. In an era of increasing climate change creating novel winter conditions, continued monitoring both long‐term and following extreme events is needed to understand new species–climate dynamics.

Environmental Conditions and Species Identity Drive Metabolite Levels in Green Leaves and Leaf Litter of 14 Temperate Woody Species

Research Highlights: Leaf chemistry is a key driver of litter decomposition; however, studies directly comparing metabolites that are important for tree growth and defence across different woody species are scarce. Background and Objectives: Choosing 14 temperate woody species differing in their growth rates, nutrient demand, shade tolerance, and drought sensitivity, we hypothesized that the species would group according to their metabolite profiles based on their ecological background. Materials and Methods: We analysed total N and C, soluble amino acid, protein, and phenolic levels in green leaves and leaf litter of these species, each in two consecutive years. Results: Metabolite levels varied significantly across species and between the sampling years which differed in temperature and precipitation (i.e., colder/drier vs warmer/ wetter). Conclusions: The 14 woody species could not be grouped according to their green leaf or leaf litter metabolite profiles. In litter leaves, most of the variation was explained by total phenolics and total nitrogen levels, and in green leaves by total phenolics and total soluble amino acid levels. Local climate variation between the two consecutive years for green leaves or leaf litter led to significant differences in metabolite levels, although some of them were species-specific.

Seasonal changes of sucrose transporter expression and sugar partitioning in common European tree species.

In temperate woody species, carbon transport from source to sink tissues is a striking physiological process, particularly considering seasonal changes. The functions of different tissues can also alternate across the seasons. In this regard, phloem loading and sugar distribution are important aspects of carbon partitioning, and sucrose uptake transporters (SUTs) play a key role in these processes. Therefore, the influence of seasons and different light-dark conditions on the expression of SUTs from 3-year-old Fagus sylvatica L., Quercus robur L. and Picea abies (L.) Karst. trees were analyzed. In addition, tissue-specific sugar and starch contents under these different environmental conditions were determined. Putative SUTs were identified in the gymnosperms (Picea abies, Ginkgo biloba L.), here for the first time, and also in the angiosperms (Q. robur, F. sylvatica). The identified SUT sequences of the different tree species cluster into three types, similar to other SUTs from herbaceous and tree species. Furthermore, the sequences from angiosperm and those from gymnosperm species form distinct clusters within the three types of SUTs. In F. sylvatica, Q. robur and P. abies, the expression levels of the different SUTs during seasons showed marked variations. Because of the high expression levels of type I SUTs in bark, wood and leaves during active growing phases in spring and summer, it can be assumed that they are involved in phloem loading, sucrose retrieval and possibly in further physiological processes. The expression patterns also indicate a flexible expression in all tissues depending on physiological requirements and environmental conditions. Compared with type I SUTs, the seasonal variations of type II SUT expression were less pronounced, whereas the seasonal variations of the type III SUT expression patterns were partly reverse. In addition to the seasonal regulation, the expressions of the different SUTs were also regulated by light in a diurnal manner.

The relative roles of local climate adaptation and phylogeny in determining leaf-out timing of temperate tree species

BackgroundLeaf out times of temperate forest trees are a prominent determinant of global carbon dynamics throughout the year. Abiotic cues of leaf emergence are well studied but investigation of the relative roles of shared evolutionary history (phylogeny) and local adaptation to climate in determining the species-level responses to these cues is needed to better apprehend the effect of global change on leaf emergence. We explored the relative importance of phylogeny and climate in determining the innate leaf out phenology across the temperate biome.MethodsWe used an extensive dataset of leaf-out dates of 1126 temperate woody species grown in eight Northern Hemisphere common gardens. For these species, information on the native climate and phylogenetic position was collected. Using linear regression analyses, we examine the relative effect of climate variables and phylogeny on leaf out variation among species.ResultsClimate variables explained twice as much variation in leaf out timing as phylogenetic information, a process that was driven primarily by the complex interactive effects of multiple climate variables. Although the primary climate factors explaining species-level variation in leaf-out timing varied drastically across different families, our analyses reveal that local adaptation plays a stronger role than common evolutionary history in determining tree phenology across the temperate biome.ConclusionsIn the long-term, the direct effects of physiological adaptation to abiotic effects of climate change on forest phenology are likely to outweigh the indirect effects mediated through changes in tree species composition.

Evolution of genome sizes in Chinese Bambusoideae (Poaceae) in relation to karyotype

DNA contents of Chinese 134 bamboo species were first time announced, and FISH of 45S rDNA shows that paleotropical woody bamboo species may be originated from a multi-source system. Woody bamboo mainly includes two types of species: the temperate woody bamboo and the tropical woody bamboo. Both vary widely in genome sizes, chromosome number, and ploidy. In this study, the DNA contents of 135 bamboo species (22 genera) mainly distributed in China were analyzed by the flow cytometry. The DNA contents of the 134 bamboo species were first time announced. In contrast to the paleotropical woody bamboos, the temperate woody species is larger in genome size. Both the average arm ratio and the average symmetric coefficient of karyotype of the temperate woody species are less than those of the paleotropical woody bamboos, suggesting that the karyotypes of temperate woody bamboo species are more symmetrical than that of paleotropical bamboos. Also, FISH (fluorescence in situ hybridization) was applied to evaluate the distribution of 45S rDNA in the chromosomes of the representative bamboo species. Only one pair of sites was detected for 45S rDNA in temperate woody bamboo species, while three pairs of sites in paleotropical woody bamboo species, implying that the paleotropical woody bamboo species may be originated from a multi-source system. This study exposed a corner of the complex genomic relationships in the woody bamboos and provided an initial framework for future phylogenic studies.

Characterization, localization, and seasonal changes of the sucrose transporter FeSUT1 in the phloem of Fraxinus excelsior.

Trees are generally assumed to be symplastic phloem loaders. A typical feature for most wooden species is an open minor vein structure with symplastic connections between mesophyll cells and phloem cells, which allow sucrose to move cell-to-cell through the plasmodesmata into the phloem. Fraxinus excelsior (Oleaceae) also translocates raffinose family oligosaccharides in addition to sucrose. Sucrose concentration was recently shown to be higher in the phloem sap than in the mesophyll cells. This suggests the involvement of apoplastic steps and the activity of sucrose transporters in addition to symplastic phloem-loading processes. In this study, the sucrose transporter FeSUT1 from F. excelsior was analysed. Heterologous expression in baker's yeast showed that FeSUT1 mediates the uptake of sucrose. Immunohistochemical analyses revealed that FeSUT1 was exclusively located in phloem cells of minor veins and in the transport phloem of F. excelsior. Further characterization identified these cells as sieve elements and possibly ordinary companion cells but not as intermediary cells. The localization and expression pattern point towards functions of FeSUT1 in phloem loading of sucrose as well as in sucrose retrieval. FeSUT1 is most likely responsible for the observed sucrose gradient between mesophyll and phloem. The elevated expression level of FeSUT1 indicated an increased apoplastic carbon export activity from the leaves during spring and late autumn. It is hypothesized that the importance of apoplastic loading is high under low-sucrose conditions and that the availability of two different phloem-loading mechanisms confers advantages for temperate woody species like F. excelsior.

Increased heat requirement for leaf flushing in temperate woody species over 1980-2012: effects of chilling, precipitation and insolation.

Recent studies have revealed large unexplained variation in heat requirement-based phenology models, resulting in large uncertainty when predicting ecosystem carbon and water balance responses to climate variability. Improving our understanding of the heat requirement for spring phenology is thus urgently needed. In this study, we estimated the species-specific heat requirement for leaf flushing of 13 temperate woody species using long-term phenological observations from Europe and North America. The species were defined as early and late flushing species according to the mean date of leaf flushing across all sites. Partial correlation analyses were applied to determine the temporal correlations between heat requirement and chilling accumulation, precipitation and insolation sum during dormancy. We found that the heat requirement for leaf flushing increased by almost 50% over the study period 1980-2012, with an average of 30 heat units per decade. This temporal increase in heat requirement was observed in all species, but was much larger for late than for early flushing species. Consistent with previous studies, we found that the heat requirement negatively correlates with chilling accumulation. Interestingly, after removing the variation induced by chilling accumulation, a predominantly positive partial correlation exists between heat requirement and precipitation sum, and a predominantly negative correlation between heat requirement and insolation sum. This suggests that besides the well-known effect of chilling, the heat requirement for leaf flushing is also influenced by precipitation and insolation sum during dormancy. However, we hypothesize that the observed precipitation and insolation effects might be artefacts attributable to the inappropriate use of air temperature in the heat requirement quantification. Rather than air temperature, meristem temperature is probably the prominent driver of the leaf flushing process, but these data are not available. Further experimental research is thus needed to verify whether insolation and precipitation sums directly affect the heat requirement for leaf flushing.

Measuring biomass and carbon stock in resprouting woody plants.

Resprouting multi-stemmed woody plants form an important component of the woody vegetation in many ecosystems, but a clear methodology for reliable measurement of their size and quick, non-destructive estimation of their woody biomass and carbon stock is lacking. Our goal was to find a minimum number of sprouts, i.e., the most easily obtainable, and sprout parameters that should be measured for accurate sprout biomass and carbon stock estimates. Using data for 5 common temperate woody species, we modelled carbon stock and sprout biomass as a function of an increasing number of sprouts in an interaction with different sprout parameters. The mean basal diameter of only two to five of the thickest sprouts and the basal diameter and DBH of the thickest sprouts per stump proved to be accurate estimators for the total sprout biomass of the individual resprouters and the populations of resprouters, respectively. Carbon stock estimates were strongly correlated with biomass estimates, but relative carbon content varied among species. Our study demonstrated that the size of the resprouters can be easily measured, and their biomass and carbon stock estimated; therefore, resprouters can be simply incorporated into studies of woody vegetation.

Chilling requirement of Ribes cultivars.

It is usually thought that adequate winter chill is required for the full flowering of many temperate woody species. This paper investigates the sensitivity of blackcurrant bud burst and flowering to natural weather fluctuations in a temperate maritime climate, and compares a range of chill models that have been proposed for assessing the accumulation of winter chill. Bud break for four contrasting cultivars are compared in an exceptionally cold and in a mild winter in Eastern Scotland. The results confirm the importance of chilling at temperatures lower than 0°C and demonstrate that no single chilling function applies equally to all blackcurrant cultivars. There is a pressing need for further model development to take into account the relationship between chilling temperatures and warming temperatures occurring both during and after the chill accumulation period.

Interspecific variation in the size‐dependent resprouting ability of temperate woody species and its adaptive significance

Summary Resprouting of woody species after above‐ground damage may help plants to persist longer at a given site and quickly reoccupy disturbed sites, thereby strongly influencing forest dynamics. Resprouting has been discussed from two adaptation perspectives: recovery from damage by catastrophic disturbance and survival in frequently disturbed shaded understorey. However, few studies have comprehensively dealt with both adaptation types to understand resprouting strategies. To understand the adaptive significance of resprouting, we assessed the size dependence of resprouting ability after stem clipping for 24 deciduous broad‐leaved species, including shrubs, sub‐canopy and canopy trees, in a cool‐temperate forest in Japan. The community assembly includes species adapted to past catastrophic disturbances (e.g. fire, logging) and to stable forest with intermittent treefall (currently the dominant disturbance). We correlated resprouting ability with life‐history strategies based on demographic parameters and plant functional traits, such as leaf mass per area (LMA), leaf toughness and wood density. All the studied species could resprout in juveniles, and resprouting ability increased as stump size increased. Most sub‐canopy and canopy trees lost their ability to resprout after attaining a particular stump size, whereas shrub species retained the ability to resprout throughout their lifetimes. The relative growth rate, LMA and foliar nitrogen did not greatly influence the resprouting ability of a species. In contrast, species with smaller maximum size, lower leaf toughness and lower wood density had better juvenile resprouting ability. This better resprouting ability may have evolved because these characteristics make them more vulnerable to shaded understorey. However, species with larger maximum size and lower leaf toughness retained their ability to resprout to a larger size. Synthesis. A better resprouting ability is related to the ability to survive frequent disturbances, in juveniles, which are characteristics of both forest understorey and frequent fire or drought. To retain resprouting ability until grown seems to be an adaptation to survive infrequent large disturbances. Light‐demanding species, which generally have better resprouting ability than shade‐tolerants both in juveniles and adults, are adapted to disturbances of various scale and frequency; however, shade‐tolerants could survive well in the understorey due to a combination of stronger physical defences and resprouting ability.

Demography of the upward-shifting temperate woody species of the Rhododendron pseudochrysanthum complex and ecologically relevant adaptive divergence in its trailing edge populations

Trailing edge populations of upward migrating species induced by postglacial climatic warming can be evolutionarily significant in the face of global warming. We tested for population size changes between lower- and higher-elevation populations located in the same mountainous regions of the upward-shifting temperate woody species of the Rhododendron pseudochrysanthum complex in Taiwan. We also tested whether natural selection evoked adaptive divergence in trailing edge populations of this species complex. We genotyped 26 expressed sequence tag-simple sequence repeat (EST-SSR) loci of 185 individuals from nine populations of the R. pseudochrysanthum complex including Rhododendron rubropunctatum, Rhododendron hyperythrum, Rhododendron morii, and R. pseudochrysanthum. R. rubropunctatum populations in the R. pseudochrysanthum complex possessed the lowest estimates of genetic diversity and effective population size. Higher-elevation R. pseudochrysanthum populations had lower effective population sizes than lower-elevation R. morii populations in Hohuanshan and Tahsueshan, as revealed by estimates using both MIGRATE-N and approximate Bayesian computation (ABC). R. rubropunctatum populations diverged significantly from populations of other members of the R. pseudochrysanthum complex. An outlier potentially under positive selection specific to R. rubropunctatum populations was identified and strongly associated with ecologically relevant environmental variables. Postglacial climatic warming has a significant influence on population isolation in the R. pseudochrysanthum complex. The results indicate adaptive evolution in the trailing edge populations, i.e., R. rubropunctatum populations of the upward shifting R. pseudochrysanthum complex.

Recovery performance in xylem hydraulic conductivity is correlated with cavitation resistance for temperate deciduous tree species

Woody species hydraulically vulnerable to xylem cavitation may experience daily xylem embolism. How such species cope with the possibility of accumulated embolism is unclear. In this study, we examined seven temperate woody species to assess the hypothesis that low cavitation resistance (high vulnerability to cavitation) is compensated by high recovery performance via vessel refilling. We also evaluated leaf functional and xylem structural traits. The xylem recovery index (XRI), defined as the ratio of xylem hydraulic conductivity in plants rewatered after soil drought to that in plants under moist conditions, varied among species. The xylem water potential causing 50% loss of hydraulic conductivity (Ψ50) varied among the species studied, whereas only a slight difference was detected with respect to midday xylem water potential (Ψmin), indicating smaller hydraulic safety margins (Ψmin - Ψ50) for species more vulnerable to cavitation. Cavitation resistance (|Ψ50|) was negatively correlated with XRI across species, with cavitation-vulnerable species showing a higher performance in xylem recovery. Wood density was positively correlated with cavitation resistance and was negatively correlated with XRI. These novel results reveal that coordination exists between cavitation resistance and xylem recovery performance, in association with wood functional traits such as denser wood for cavitation-resistant xylem and less-dense but water-storable wood for refillable xylem. These findings provide insights into long-term maintenance of water transport in tree species growing under variable environmental conditions.

Testing the packing rule across the twig–petiole interface of temperate woody species

Theories that incorporate the area-preserving rule of Leonardo da Vinci predict that the sum of the cross-sectional lumen area of xylem conduits (vessels or tracheids) is constant across different levels of branching. If true, this rule obtains the packing rule, according to which (1) vessel cross-sectional lumen area (A) will negatively and isometrically scale to vessel number per unit area (N) and (2) the distal to proximal vessel diameter ratio (DR = D (n+1)/D (n) ) should scale as the -1/2 power of the distal to proximal vessel number ratio (NR = N (n+1)/N (n) ), i.e., DR = NR-1/2. Using data collected from the terminal twigs and the petioles of 60 temperate (27 evergreen and 33 deciduous) woody species from southwestern China, we determined the scaling relationships for A versus N and for DR versus NR. Analyses of the data revealed contrasting scaling exponents and normalization constants for A versus N and for DR versus NR and no consistent trend across the two species groupings or between the two organ types (as predicted by the area-preserving and packing rules). These results caution against applying these rules ubiquitously to all species or to different organ types (even on the same plant).

Modeling the influence of differential sectoriality on the photosynthetic responses of understory saplings to patchy light and water availability

Exploitation of patchy light is a key determinant of plant performance in the forest understory. While many adaptive traits are known, the role of stem vasculature in understory photosynthesis is not established. Sectoriality—the degree of vascular constraint to long distance transport—has been hypothesized to limit growth in heterogeneous light. We simulated the photosynthetic potential of sectored and integrated plants in patchy light, as a function of soil water potential (patchy or uniform). We used hydraulic parameters typical of temperate woody species in an Ohm’s law model including a tangential resistance parameter, and simulated cavitation by varying axial resistance of leaves, leaves and roots, or the whole plant. Our results suggest that differential sectoriality will not affect photosynthesis when water is plentiful, but can constrain stomatal conductance at more negative soil water potentials, especially when only a small portion of the crown receives light. This effect is strongest just below the turgor loss point, and depends on axial resistance and soil water heterogeneity. Increased resistance in high light leaves decreases photosynthesis regardless of sectoriality. However, when resistance is increased for leaves and roots or the whole plant, photosynthesis decreases more for sectored than for integrated plants. Moreover, the simulations suggest that sectoriality can further depress photosynthesis when water availability is asymmetrical. These results might explain why integrated species, such as Betula lenta, B. alleghaniensis, and Acer saccharum thrive in the forest understory and grow rapidly into canopy gaps, while sectored species, such as Quercus rubra, do not.

The productivity–diversity relationship: varying aims and approaches

The productivity-diversity relationship is an impor tant and heavily debated issue in ecology. A major advancement in exploring this relationship has been the analyses of large numbers of case-studies (Mittelbach et al. 2001, Gillman and Wright 2006). These works have shown that the relationship between productivity and diversity varies, and that a single general relationship does not exist. We investigated the productivity diversity relationship with respect to the species pool concept (P?rtel et al. 2007), which postulates that more species are expected to evolve in conditions (ecosystems) that have been historically more common. Tropical humid ecosystems have been relatively more productive: during the past hundreds of millions of years, whereas productivity in temperate ecosystems has been limited by low temperatures and repeated glaciations. We found that positive productivity-diversity relationships are more common in the tropics, where the species pool of productive habitats is large, whereas declines in species richness at high productivities is more common in temperate regions, since species pools in high produc tivity ecosystems are likely to be relatively small. Similarly, we showed that positive productivity-diversi ty relationship can be common even in temperate regions, if only woody species diversity is considered (Laanisto et al. 2008). This finding might reflect evolutionary history: most temperate woody species originate from lineages and exhibit tropical patterns due to niche conservatism. Robert J. Whittaker (2010, hereafter RJW) criticizes our above-mentioned studies, claiming we were too liberal in selecting case studies. RJW's argument relies on the assumption that the seven conditions for case studies he describes as reasonable and necessary criteria, are always valid. We argue that different research aims require a priori criteria to be defined. Demanding implementation of criteria as hindsight criticism of existing studies can easily lead to paradoxes (e.g., richness rather than diversity should be used for diversity relationships). Our approach in selecting case studies for analysis was straightforward: a case study must reveal a suitable habitat-productivity-plant-diver

Alteration of PHYA expression change circadian rhythms and timing of bud set in Populus

In many temperate woody species, dormancy is induced by short photoperiods. Earlier studies have shown that the photoreceptor phytochrome A (phyA) promotes growth. Specifically, Populus plants that over-express the oat PHYA gene (oatPHYAox) show daylength-independent growth and do not become dormant. However, we show that oatPHYAox plants could be induced to set bud and become cold hardy by exposure to a shorter, non-24 h diurnal cycle that significantly alters the relative position between endogenous rhythms and perceived light/dark cycles. Furthermore, we describe studies in which the expression of endogenous Populus tremula x P. tremuloides PHYTOCHROME A (PttPHYA) was reduced in Populus trees by antisense inhibition. The antisense plants showed altered photoperiodic requirements, resulting in earlier growth cessation and bud formation in response to daylength shortening, an effect that was explained by an altered innate period that leads to phase changes of clock-associated genes such as PttCO2. Moreover, gene expression studies following far-red light pulses show a phyA-mediated repression of PttLHY1 and an induction of PttFKF1 and PttFT. We conclude that the level of PttPHYA expression strongly influences seasonally regulated growth in Populus and is central to co-ordination between internal clock-regulated rhythms and external light/dark cycles through its dual effect on the pace of clock rhythms and in light signaling.