Leaf size serves as a proxy for xylem vulnerability to cavitation in plantation trees

Hybrid poplars are an important renewable forest resource known for their high productivity. At the same time, they are highly vulnerable to water stress. Identifying traits that can serve as indicators for growth performance remains an important task, particularly under field conditions. Understanding which trait combinations translate to improved productivity is key in order to satisfy the demand for poplar wood in an uncertain future climate. In this study, we compared hydraulic and leaf traits among five hybrid poplar clones at 10 plantations in central Alberta. We also assessed the variation of these traits between 2‐ to 3‐year‐old branches from the lower to mid‐crown and current‐year long shoots from the mid to upper crown. Our results showed that (1) hybrid poplars differed in key hydraulic parameters between branch type, (2) variation of hydraulic traits among clones was relatively large for some clones and less for others, and (3) strong relationships between measured hydraulic traits, such as vessel diameter, cavitation resistance, xylem‐specific and leaf‐specific conductivity and leaf area, were observed. Our results suggest that leaf size could serve as an additional screening tool when selecting for drought‐tolerant genotypes in forest management and tree improvement programmes.     

Mechanisms underlying the effects of fall dormancy on the cold acclimation and winter hard- iness of Medicago sativa

As a global planting forage legume, alfalfa (Medicago sativa) is a valuable material to study the evolutionary and ecological mechanisms on plant adaptation to freezing due to their contrasting winter hardiness induced by fall dormancy (FD). This paper reveals that FD is an important growth characteristic that is adaptative to short-day and reducing temperature in late autumn, followed by a higher overwintering rate. Alfalfa cultivars are grouped into 11 FD ratings (numbered from 1 to 11), and this phenomenon is related to the extensive spread and cultivation for thousands of years in the globe. Alfalfa cultivars are under different climate habitats for a long time, leading to FD adaptive evolution, which provides rich genetic resources for human. In general, adaptative process associated with cold acclimation and winter hardiness in alfalfa is affected by FD, thus differences in winter hardiness exist among alfalfa cultivars. So far, regulation of FD by light and temperature and effects of FD on physiological and ecological processes involved in cold acclimation have been reported largely. However, signal transduction and the regulatory network associated with gene expression, especially the molecular mechanisms by which antifreeze proteins function in cold adaptation, are still poorly understood. Several scientific problems that need to be addressed in the future studies are highlighted in this review.     

紫花苜蓿秋眠性对低温驯化过程与越冬耐寒适应的作用机理

紫花苜蓿(Medicago sativa)是全球性的栽培牧草, 因其秋眠性强弱而导致的抗寒性差异为深刻解析植物耐寒适应的进化生态学机制提供了一个很好的研究模式。作为多年生豆科植物, 秋眠性是紫花苜蓿适应晚秋日照缩短、温度降低, 以及提高越冬存活率的一种生长特性。秋眠性的形成与几千年来紫花苜蓿在全球的传播扩展和栽培利用有关, 长期处于不同的气候生境, 导致秋眠性的适应进化, 这为人类利用提供了丰富的遗传资源。根据秋眠性的强弱, 学术界目前将之分为11个等级。一般而言, 秋眠性的强弱影响了紫花苜蓿的低温驯化与越冬耐寒适应等过程, 导致不同品种间的抗寒性存在差异。迄今, 关于秋眠性的光温调控, 以及秋眠性影响低温驯化的生理生态过程研究较多, 而对相关的细胞信号转导与基因表达调控途径, 尤其是对秋眠性如何影响越冬紫花苜蓿抗冻蛋白作用等耐寒适应的分子机制, 尚知之甚少。针对目前研究中存在的问题, 该文提出了未来需要重点关注的科学问题。     

Low night temperature and inhibition of gibberellin biosynthesis override phytochrome action and induce bud set and cold acclimation, but not dormancy in PHYA overexpressors and wild-type of hybrid aspen

Juvenile trees of temperate and boreal regions cease growth and set buds in autumn in response to short day-lengths (SD) detected by phytochrome. Growth cessation and bud set are prerequisites for the development of winter dormancy and full cold hardiness. In this study we show that the SD-requirement for bud set and cold hardening can be overcome in hybrid aspen (Populus tremula L. × tremuloides Michx.) by low night temperature and inhibition of gibberellin (GA) biosynthesis. Bud set and increased cold hardiness were observed under normally non-inductive long day-length (LD) in wild-type plants, when exposed to low night temperature and paclobutrazol. In addition, the effect of PHYA overexpression could be overcome in transgenic plants, producing bud set and cold acclimation by treatment with: SD, low night temperature and paclobutrazol. After cold acclimation, the degree of bud dormancy was lower for wild-type plants prior treated with LD and transgenic plants (overexpressing PHYA), than SD-treated, wild-type plants. Thus, low night temperature in combination with reduced GA content induced bud set and promoted cold hardiness under normally non-inductive photoperiods in hybrid aspen, but was unable to affect development of dormancy. This might suggest separate signalling pathways from phytochrome regulating the induction of cold/cold hardiness and bud dormancy in hybrid aspen or alternatively, there might be one pathway that fails to complete its action in the transgenic and paclobutrazol treated plants.     

Causes of variation in cold hardiness among fast-growing willows (Salix spp.) with particular reference to their inherent rates of cold hardening

Causes of variation in cold hardiness in the autumn were assessed among closely related, fast‐growing clones of willow of northern/continental and southern/maritime origins, under controlled regimes and natural conditions. Cold hardiness was assessed by controlled freezing followed by injury analysis, based on measurements of chlorophyll fluorescence (stems) and electrolyte leakage (leaves). During growth at a given temperature, the cold hardiness of the clones' stems was negatively correlated with their rate of growth. This apparently phenotypic variation was independent of temperature and, hence, the absolute growth rate. At later stages, cold hardiness of stems varied mainly with respect to genetic differences in the timing and rate of cold hardening. Cold hardening began up to 7 weeks earlier in northern/continental clones, and their rates of hardening in cool temperature regimes were up to three times higher than in southern/maritime clones. Ranking of clones with respect to rates was essentially the same whether natural or abrupt reductions of day length were used to trigger cold hardening. Results closely agreed with those of a previous field trial. Comparisons of rates at cool and warm temperatures suggest that cold hardening became increasingly dependent on cool temperatures with time. Increasing sucrose‐to‐glucose ratios, and especially dry‐to‐fresh weight ratios, paralleled early cold hardening. Before leaves were shed in the autumn they underwent cold hardening in parallel with stems, eventually allowing them to tolerate temperatures down to ?10 °C.     

Ecotype-dependent control of growth,dormancy and freezing tolerance under seasonal changes in <Emphasis Type="Italic">Betula pendula</Emphasis> Roth

Woody plants in the temperate and boreal zone undergo annual cycle of growth and dormancy under seasonal changes. Growth cessation and dormancy induction in autumn are prerequisites for the development of substantial cold hardiness in winter. During evolution, woody plants have developed different ecotypes that are closely adapted to the local climatic conditions. In this study, we employed distinct photoperiodic ecotypes of silver birch (Betula pendula Roth) to elucidate differences in these adaptive responses under seasonal changes. In all ecotypes, short day photoperiod (SD) initiated growth cessation and dormancy development, and induced cold acclimation. Subsequent low temperature (LT) exposure significantly enhanced freezing tolerance but removed bud dormancy. Our results suggested that dormancy and freezing tolerance might partially overlap under SD, but these two processes were regulated by different mechanisms and pathways under LT. Endogenous abscisic acid (ABA) levels were also altered under seasonal changes; the ABA level was low during the growing season, then increased in autumn, and decreased in winter. Compared with the southern ecotype, the northern ecotype was more responsive to seasonal changes, resulting in earlier growth cessation, cold acclimation and dormancy development in autumn, higher freezing tolerance and faster dormancy release in winter, and earlier bud flush and growth initiation in spring. In addition, although there was no significant ecotypic difference in ABA level during growing season, the rates and degrees of ABA alterations were different between the ecotypes in autumn and winter, and could be related to ecotypic differences in dormancy and freezing tolerance.     

Use of intraspecific variation in thermal responses for estimating an elevational cline in the timing of cold hardening in a sub‐boreal conifer

To avoid winter frost damage, evergreen coniferous species develop cold hardiness with suitable phenology for the local climate regime. Along the elevational gradient, a genetic cline in autumn phenology is often recognised among coniferous populations, but further quantification of evolutionary adaptation related to the local environment and its responsible signals generating the phenological variation are poorly understood. We evaluated the timing of cold hardening among populations of Abies sachalinensis, based on time series freezing tests using trees derived from four seed source populations × three planting sites. Furthermore, we constructed a model to estimate the development of hardening from field temperatures and the intraspecific variations occurring during this process. An elevational cline was detected such that high‐elevation populations developed cold hardiness earlier than low‐elevation populations, representing significant genetic control. Because development occurred earlier at high‐elevation planting sites, the genetic trend across elevation overlapped with the environmental trend. Based on the trade‐off between later hardening to lengthen the active growth period and earlier hardening to avoid frost damage, this genetic cline would be adaptive to the local climate. Our modelling approach estimated intraspecific variation in two model components: the threshold temperature, which was the criterion for determining whether the trees accumulated the thermal value, and the chilling requirement for trees to achieve adequate cold hardiness. A higher threshold temperature and a lower chilling requirement could be responsible for the earlier phenology of the high‐elevation population. These thermal responses may be one of the important factors driving the elevation‐dependent adaptation of A. sachalinensis.     

Drought tolerance and growth in populations of a wide‐ranging tree species indicate climate change risks for the boreal north

Choosing drought‐tolerant planting stock in reforestation programs may help adapt forests to climate change. To inform such reforestation strategies, we test lodgepole pine (Pinus contorta Doug. ex Loud. var latifolia Englm.) population response to drought and infer potential benefits of a northward transfer of seeds from drier, southern environments. The objective is addressed by combining dendroecological growth analysis with long‐term genetic field trials. Over 500 trees originating from 23 populations across western North America were destructively sampled in three experimental sites in southern British Columbia, representing a climate warming scenario. Growth after 32 years from provenances transferred southward or northward over long distances was significantly lower than growth of local populations. All populations were affected by a severe natural drought event in 2002. The provenances from the most southern locations showed the highest drought tolerance but low productivity. Local provenances were productive and drought tolerant. Provenances from the boreal north showed lower productivity and less drought tolerance on southern test sites than all other sources, implying that maladaptation to drought may prevent boreal populations from taking full advantage of more favorable growing conditions under projected climate change.     

Variable increases in cold hardiness induced in winter rape by plant growth regulators

Triazole and conventional growth regulators were tested for their ability to extend cold hardiness and improve the winter survival of winter rape (Brassica napus L.). Winter rape plants were grown in the field (Ottawa 45°23 N) and in growth cabinets. Plant growth regulators (PGRs) were applied during the early vegetative stage and the plants were allowed to cold harden. Cold-hardened plants from the field and cabinet were subjected to freezing and ice encasement tests in the laboratory. Some of the triazole PGRs reduced plant size by limiting cell expansion and increasing cell numbers. While cold hardiness and ice encasement tolerance were increased by some growth regulators, these effects were not consistent with time nor were they reflected in increased winter survival. Natural cold hardening may have eclipsed the PGR-induced hardening.PGR contribution number 1381.     

Effect of photoperiod and temperature on the development of frost hardiness in three Alnus species

The influence of short day and low temperature on cold acclimation of A. crispa (Ait.) Pursh, A. glutinosa (L.) Gaertn. and A. rubra Bong, was investigated. Two clones of each species originating from in vitro propagation were exposed to three daylength/temperature treatments. Periodically plantlets were exposed to controlled freezing temperature in order to evaluate their level of frost hardiness.
Short day (SD) and cold temperature (CT) and long day (LD) and cold temperature (CT) were the most effective treatments for the development of frost hardiness in shoots and roots of the three species tested. Short day (SD) and warm temperature (WT) induced a significant increase in hardiness in shoots of all three species. However, this treatment did not trigger root hardening. A. crispa was found to be the hardiest species followed by A. glutinosa and A. rubra . Intraspecific variation was observed between the two A. glutinosa clones. A glutinosa clone AG8, a Russian provenance, showed a greater freezing resistance than A. glutinosa clone AG2, a German provenance.     

Changes in carbohydrates, ABA and bark proteins during seasonal cold acclimation and deacclimation in Hydrangea species differing in cold hardiness

Cold injury is frequently seen in the commercially important shrub Hydrangea macrophylla but not in Hydrangea paniculata. Cold acclimation and deacclimation and associated physiological adaptations were investigated from late September 2006 to early May 2007 in stems of field-grown H. macrophylla ssp. macrophylla (Thunb.) Ser. cv. Blaumeise and H. paniculata Sieb. cv. Kyushu. Acclimation and deacclimation appeared approximately synchronized in the two species, but they differed significantly in levels of mid-winter cold hardiness, rates of acclimation and deacclimation and physiological traits conferring tolerance to freezing conditions. Accumulation patterns of sucrose and raffinose in stems paralleled fluctuations in cold hardiness in both species, but H. macrophylla additionally accumulated glucose and fructose during winter, indicating species-specific differences in carbohydrate metabolism. Protein profiles differed between H. macrophylla and H. paniculata, but distinct seasonal patterns associated with winter acclimation were observed in both species. In H. paniculata concurrent increases in xylem sap abscisic acid (ABA) concentrations ([ABA](xylem)) and freezing tolerance suggests an involvement of ABA in cold acclimation. In contrast, ABA from the root system was seemingly not involved in cold acclimation in H. macrophylla, suggesting that species-specific differences in cold hardiness may be related to differences in [ABA](xylem). In both species a significant increase in stem freezing tolerance appeared long after growth ceased, suggesting that cold acclimation is more regulated by temperature than by photoperiod.     

木本植物木质部的冻融栓塞应对研究进展

冻融栓塞在中高纬度地区木本植物中普遍存在。抗冻融栓塞能力对在寒冷环境中木本植物的生长和安全越冬十分关键, 这直接决定植物分布范围。冻融栓塞是由于冰中气体溶解度低, 木质部水分在低温下冷冻, 使之前水中溶解的气体逸出到导管中, 随后木质部中的冰融化又使气泡扩张而引发的栓塞现象。木质部解剖结构的差异会影响植物的抗冻融栓塞能力, 植物还可以通过调节木质部正压、代谢耗能等方式主动修复冻融栓塞, 也可通过增加树液溶质含量等逃避冷冻, 以减少低温损伤。然而, 与干旱栓塞相比, 目前对木质部冻融栓塞的形成以及植物响应和调节机制的理解不足。为此, 该文首先综述了木质部冻融栓塞的形成机制和植物的逃避、忍耐、修复等3种冻融栓塞的应对策略, 然后总结了木质部抗低温胁迫能力的生理表现、影响因子和评价指标, 并在此基础上讨论了低温抗性、干旱抗性和水力效率之间的多元权衡关系, 最后提出今后该领域中的5个优先研究问题: (1)不同植物冰冻的最低温度阈值; (2)是否存在应对低温胁迫的水力脆弱性分割机制; (3)冻融栓塞修复与代谢消耗的关系; (4)低温抗性、干旱抗性和水力效率之间的权衡关系; (5)抗冻融栓塞性状是否能够纳入经济性状谱系。     

Factors affecting the cold hardiness of the peach-potato aphid Myzus persicae

Supercooling point studies were used to investigate the factors influencing the cold hardiness of the peach-potato aphid Myzus persicae, a freezing-susceptible insect. Overwintering adults lost cold hardiness as winter progressed, with a variable proportion showing a marked reduction in supercooling ability. Cold hardiness increased in spring so that all individuals demonstrated extensive supercooling ability typical of aphids reared in the laboratory at 20°C with a long photoperiod; these levels of cold hardiness were maintained in the field during summer and early autumn. First instar nymphs demonstrated considerable cold hardiness all year. Surface moisture caused inoculative freezing in some first instar nymphs and adults when supercooled, but the majority were unaffected. In the laboratory, adults starved for 7 days at 5°C showed distinct losses of supercooling potential equivalent to those observed in the field during mid to late winter. No loss of cold hardiness was found in first instar nymphs starved under the same conditions. The results demonstrate that the cold hardiness characteristics of M. persicae are atypical of those observed in other freezing-susceptible insects and it is suggested that continued feeding during mild winter conditions allows maintenance of cold hardiness particularly in adult aphids, and provides a possible explanation for the successful anholocyclic overwintering of M. persicae during such winters.     

Ice segregation in the crown of winter cereals: Evidence for extraorgan and extratissue freezing

Meaningful improvements in winter cereal cold hardiness requires a complete model of freezing behaviour in the critical crown organ. Magnetic resonance microimaging diffusion‐weighted experiments provided evidence that cold acclimation decreased water content and mobility in the vascular transition zone (VTZ) and the intermediate zone in rye (Secale cereale L. Hazlet) compared with wheat (Triticum aestivum L. Norstar). Differential thermal analysis, ice nucleation, and localization studies identified three distinct exothermic events. A high‐temperature exotherm (?3°C to ?5°C) corresponded with ice formation and high ice‐nucleating activity in the leaf sheath encapsulating the crown. A midtemperature exotherm (?6°C and ?8°C) corresponded with cavity ice formation in the VTZ but an absence of ice in the shoot apical meristem (SAM). A low‐temperature exotherm corresponded with SAM injury and the killing temperature in wheat (?21°C) and rye (?27°C). The SAM had lower ice‐nucleating activity and freezing survival compared with the VTZ when frozen in vitro. The intermediate zone was hypothesized to act as a barrier to ice growth into the SAM. Higher cold hardiness of rye compared with wheat was associated with higher VTZ and intermediate zone desiccation resulting in the formation of ice barriers surrounding the SAM.     

Agronomic performance of 27 Populus clones evaluated after two 3‐year coppice rotations in Henan,China

Selecting superior clones is the first step for commercial short‐rotation coppice cultures to provide biomass and bioenergy. Till date, such selection for hybrid Populus clones in middle China is absent. Here we describe the growth, aboveground biomass production and cell wall composition of 27 hybrid poplar clones in Henan, China for two 3‐year rotations. Significant variation in these three characteristics over two triennial rotation coppices among the 27 poplar clones was observed. During two 3‐year rotation coppices, clones ‘276’, ‘02‐17’, and ‘599’ showed relatively higher tree heights and larger basal diameters than those of the other clones. However, the most productive clones were ‘36’ and ‘01‐30’. At the end of the second triennial rotation, the aboveground biomass production reached 18 Mg ha^?1 year^?1. For the cell wall composition analysis, the cellulose contents of clones ‘01‐243’ and ‘2001’ were relatively high, while the xylose contents of clones ‘01‐30’ and ‘65’ were relatively high. Cluster analysis based on height, basal diameter, biomass, heat value, cellulose content, and survival rate revealed five growth potential classes. Accordingly, clones ‘03‐332’, ‘36’, and ‘599’ exhibited high biomass and growth and had the greatest potential to serve as excellent biomass producers in Henan, China. In addition, the expression patterns of 20 key regulatory genes were analyzed, and an integrated coexpression network was constructed. This field trial provides a comprehensive quantification and evaluation of the agronomic performance of 27 poplar clones in Henan, China. The results of this study and the analytical strategies provide an efficient mechanism for selecting clones that will perform well agronomically in local environments. The expression of key genes and the integrated coexpression network provide the molecular mechanisms of poplar biomass performance.     

Impact of climate change on cold hardiness of Douglas‐fir (Pseudotsuga menziesii): environmental and genetic considerations

The success of conifers over much of the world's terrestrial surface is largely attributable to their tolerance to cold stress (i.e., cold hardiness). Due to an increase in climate variability, climate change may reduce conifer cold hardiness, which in turn could impact ecosystem functioning and productivity in conifer‐dominated forests. The expression of cold hardiness is a product of environmental cues (E), genetic differentiation (G), and their interaction (G × E), although few studies have considered all components together. To better understand and manage for the impacts of climate change on conifer cold hardiness, we conducted a common garden experiment replicated in three test environments (cool, moderate, and warm) using 35 populations of coast Douglas‐fir (Pseudotsuga menziesii var. menziesii) to test the hypotheses: (i) cool‐temperature cues in fall are necessary to trigger cold hardening, (ii) there is large genetic variation among populations in cold hardiness that can be predicted from seed‐source climate variables, (iii) observed differences among populations in cold hardiness in situ are dependent on effective environmental cues, and (iv) movement of seed sources from warmer to cooler climates will increase risk to cold injury. During fall 2012, we visually assessed cold damage of bud, needle, and stem tissues following artificial freeze tests. Cool‐temperature cues (e.g., degree hours below 2 °C) at the test sites were associated with cold hardening, which were minimal at the moderate test site owing to mild fall temperatures. Populations differed 3‐fold in cold hardiness, with winter minimum temperatures and fall frost dates as strong seed‐source climate predictors of cold hardiness, and with summer temperatures and aridity as secondary predictors. Seed‐source movement resulted in only modest increases in cold damage. Our findings indicate that increased fall temperatures delay cold hardening, warmer/drier summers confer a degree of cold hardiness, and seed‐source movement from warmer to cooler climates may be a viable option for adapting coniferous forest to future climate.     

中国杨树人工林培育技术研究进展

我国杨树人工林总面积达700万hm²,位居世界第一.发展杨树人工林、满足社会经济发展和环境保护的需要是世界杨树研究的发展趋势.在介绍中国杨树栽培区区划、杨树人工林的主要栽培无性系及其生产力的基础上,总结了近10年来我国杨树在立地质量评价、苗木培育技术、人工林定向培育模式、混交林营造技术、林农复合经营技术、人工林水分和养分管理技术、立地生产力维护和杨树人工林的生态功能等方面的研究进展,讨论了我国杨树人工林的生产潜力、杨树人工林的布局、杨树人工林定向培育及加强杨树人工林的环境功能研究等问题,为我国杨树人工林资源的培育及可持续经营提供参考.     

Identification of the main site factors and management intensity affecting the establishment of Short-Rotation-Coppices (SRC) in Northern Italy through stepwise regression analysis

Data collected from 183 poplar and 102 willow SRC experimental plots, located in Central-North Italy, were subjected to stepwise regression analysis to acquire information on the environmental factors affecting plant survival and productivity in the first two-year rotation cycle. Nine Populus ×canadensis Mönch, eight P. deltoids Bartr. clones and four hybrids of Sali× matsudana Koidz were included in analysis. Independent variables were: annual and seasonal water availability (rainfall and irrigation), annual mean air temperature, soil texture, pH, N and organic matter content, planting density and management intensity. Dependent variables were: a) mean annual yield during the first two-year rotation cycle in tons per hectare per year of dry matter (Odt·ha^?1·y^?1); b) plant survival at the end of the second year from planting (%). Water availability resulted the main variable driving plant survival and biomass production in both poplar and willow clones. Water availability appeared to be the principal factor affecting the establishment of poplar and willow energy plantations in the Po valley. Possible variations in the rainfall regime consequent to climate changes could seriously influence land suitability to SRC. Experimental data also indicate that choice of planting density may increase the biomass yield during the first two-year especially with P. deltoides clones.