Xylem traits mediate a trade-off between resistance to freeze-thaw-induced embolism and photosynthetic capacity in overwintering evergreens

Hydraulic traits were studied in temperate, woody evergreens in a high-elevation heath community to test for trade-offs between the delivery of water to canopies at rates sufficient to sustain photosynthesis and protection against disruption to vascular transport caused by freeze-thaw-induced embolism. Freeze-thaw-induced loss in hydraulic conductivity was studied in relation to xylem anatomy, leaf- and sapwood-specific hydraulic conductivity and gas exchange characteristics of leaves. We found evidence that a trade-off between xylem transport capacity and safety from freeze-thaw-induced embolism affects photosynthetic activity in overwintering evergreens. The mean hydraulically weighted xylem vessel diameter and sapwood-specific conductivity correlated with susceptibility to freeze-thaw-induced embolism. There was also a strong correlation of hydraulic supply and demand across species; interspecific differences in stomatal conductance and CO(2) assimilation rates were correlated linearly with sapwood- and leaf-specific hydraulic conductivity. Xylem vessel anatomy mediated an apparent trade-off between resistance to freeze-thaw-induced embolism and hydraulic and photosynthetic capacity during the winter. These results point to a new role for xylem functional traits in determining the degree to which species can maintain photosynthetic carbon gain despite freezing events and cold winter temperatures.     

Mechanisms for tolerating freeze-thaw stress of two evergreen chaparral species: Rhus ovata and Malosma laurina (Anacardiaceae)

The response to freeze-thaw stress was examined for two co-occurring evergreen species, Malosma laurina and Rhus ovata. Laboratory and field experiments on adults and seedlings were made in the spring and winter in 1996 and again on adults in 2003 and 2004. Laboratory and field results indicated that the stem xylem for adults of M. laurina and R. ovata were similarly susceptible to freezing-induced cavitation (percentage loss of conductivity = 92 ± 2.6% for R. ovata and 90 ± 4.2% for M. laurina at ≤ -6°C). In contrast, leaves of M. laurina were more susceptible to freezing injury than leaves of R. ovata. Among seedlings in the field, leaves of M. laurina exhibited freezing injury at -4°C and total shoot mortality at -7.2°C, whereas co-occurring seedlings of R. ovata were uninjured. Surprisingly, R. ovata tolerates high levels of freezing-induced xylem embolism in the field, an apparently rare condition among evergreen plants. Rhus ovata avoids desiccation when xylem embolism is high by exhibiting low minimum leaf conductance compared to M. laurina. These results suggest a link between minimum leaf conductance and stem hydraulics as a mechanism permitting the persistence of an evergreen leaf habit in freezing environments.     

Different vulnerabilities of Quercus ilex L. to freeze- and summer drought-induced xylem embolism: an ecological interpretation

Quercus ilex L. growing in the southern Mediterranean Basin region is exposed to xylem embolism induced by both winter freezing and summer drought. The distribution of the species in Sicily could be explained in terms of the different vulnerability to embolism of its xylem conduits. Naturally occurring climatic conditions were simulated by: (1) maintaining plants for 3h at ambient temperatures of 0, -1.5, -2.5, -5.0 and -11°C; and (2) allowing plants to dry out to ratios of their minimum diurnal leaf water potentials (Ψ₁) to that at the turgor loss point (Ψ_tlp) of 0.6, 0.9, 1.05, 1.20 and 1.33. The loss of hydraulic conductivity of one-year-old twigs reached 40% at -1.5°C and at Ψ₁/Ψ_tlP= 1.05. Recovery from these strains was almost complete 24 h after the release of thermal stress or after one irrigation, respectively. More severe stresses reduced recovery consistently. The percentages of xylem conduits embolized following application of the two stresses, were positively related to xylem conduit diameter. The capability of the xylem conduits to recover from stress was positively related to the conduit diameter in plants subjected to summer drought, but not in the plants subjected to winter freezing stress. The ecological significance of the different vulnerabilities to embolism of xylem conduits under naturally occurring climatic conditions is discussed.     

Spring patterns of freezing resistance and photosynthesis of two leaf phenotypes of Hedera helix

Subdominant evergreen broad-leaved plants occurring in deciduous forests throughout temperate zones have only a short window of optimum photoassimilation in spring before canopy closure. Yet increasing photosynthetic and metabolic activity occurs concurrently with reductions in freezing resistance, resulting in vulnerability of plant tissues to late spring freezing events. Our goal was to document the temporal patterns of photosynthesis versus freezing resistance during spring in adult and juvenile leaf phenotypes of Hedera helix in Switzerland. Freezing resistances in all leaves were well below long-term minimum temperatures experienced at the study site, with adult leaf phenotypes in the forest canopy being more freezing resistant than juvenile leaves occurring closer to the ground. Reductions in freezing resistance were followed by increases in leaf photosynthetic capacities, which appeared synchronized among leaf phenotypes. Adult canopy leaves maintained a higher freezing resistance but lower photosynthetic capacity than juvenile leaves through the end of winter and into early spring. However, shortly after the cessation of freezing temperatures, adult leaves greatly increased their photosynthetic capacity relative to juvenile leaves, yet maintained freezing resistances sufficient to resist late spring freezing events. These patterns highlight the importance of the tradeoff in H. helix between exposure to potentially damaging cold temperatures in late spring and the need for high photosynthetic carbon gains before full canopy closure.     

The vulnerability to freezing-induced xylem cavitation of Larrea tridentata (Zygophyllaceae) in the Chihuahuan desert

The temperature dependence of freezing-induced xylem cavitation was studied in a Chihuahuan desert population of Larrea tridentata (Zygophyllaceae). Field measurements of wood temperature and xylem embolism were combined with anatomical studies and laboratory measurements of embolism in stem and root samples frozen under controlled conditions. Our laboratory experiments corroborated the previously observed relationship between minimum freezing temperature and embolism. The area of the low-temperature exotherms produced during the freezing treatments was correlated with the resulting embolism, suggesting that the freezing of water inside parenchyma cells is associated with the occurrence of xylem embolism. In the laboratory experiments, embolism in stems increased only at temperatures below -14°C. Although this meant that the studied population was more resistant to freezing-induced xylem embolism than a previously studied population from the Sonoran desert, the impact of freezing was nevertheless greater because of much lower environmental temperatures. This result suggests that dieback associated with periodic extreme freezes may contribute to limiting the present distribution of L. tridentata in central New Mexico. Although laboratory experiments showed that root xylem embolism increased after freezing to less negative minimum temperatures than stems (significant effects at T = -7°C), root embolism in the field was lower than shoot embolism in accordance with measured soil temperatures throughout the study.     

Acorn production is linked to secondary growth but not to declining carbohydrate concentrations in current-year shoots of two oak species

In trees, reproduction constitutes an important resource investment which may compete with growth for resources. However, detailed analyses on how growth and fruit production interact at the shoot level are scarce. Primary canopy growth depends on the development of current-year shoots and their secondary growth might also influence the number and size of fruits supported by them. We hypothesise that an enhanced thickening of current-year shoots is linked positively to acorn production in oaks. We analysed the effect of acorn production on shoot growth of two co-occurring Mediterranean oak species with contrasting leaf habit (Quercus ilex, Quercus faginea). Length and cross-sectional area of current-year shoots, apical bud mass, number of leaves and acorns, xylem and conductive area, number of vessels of acorn-bearing and non-bearing shoots were measured in summer and autumn. Nitrogen and carbohydrates analyses were also performed in stems and leaves of both shoot types. Stem cross-sectional area increased in acorn-bearing shoots when compared with non-bearing shoots for both species and such surplus secondary growth was observed since summer. In bearing shoots, the total transversal area occupied by vessels decreased significantly from basal to apical positions along the stem as did the xylem area and the number of vessels. Leaves of bearing shoots showed lower nitrogen concentration than those of non-bearing shoots. Carbohydrate concentrations did not differ in stems and leaves as a function of the presence of acorns. Such results suggest that carbohydrates may preferentially be allocated towards reproductive shoots, possibly through enhanced secondary growth, satisfying all their carbon demands for growth and reproduction. Our findings indicate that acorn production in the two studied oaks depends on shoot secondary growth.     

Differential and interactive effects of temperature and photoperiod on budburst and carbon reserves in two co-occurring Mediterranean oaks

Effects of temperature and photoperiod and their interactions on budburst and on the use of carbon reserves were examined in two Mediterranean oaks differing in wood anatomy and leaf habit. Seedlings of Quercus ilex subsp. ballota (evergreen and diffuse-porous wood) and Q. faginea (semi-deciduous and ring-porous wood) were grown under two temperatures (12 and 19 °C) and two photoperiods (10 and 16 h) in a factorial experiment. In the 16 h photoperiod at 19 °C, photosynthesis was suppressed in half of the seedlings by covering leaves with aluminium foil. The concentration of soluble sugars, starch and lipids in leaves, stems and roots was assessed before and after budburst. Under the 12 °C treatment (mean current temperature in early spring in the Iberian Peninsula), budburst in Q. faginea occurred earlier than in Q. ilex . Higher temperature promoted earlier budburst in both species, mostly under the 16 h photoperiod. This response was less pronounced in Q. faginea because its budburst was also controlled by photoperiod, and because this species needs to construct a new ring of xylem before budburst to supply its growth demands. Therefore, dates of budburst of the two species became closer to each other in the warmer treatment, which might alter competitive relations between the species with changing climate. While Q. ilex relied on carbon reserves for budburst, Q. faginea relied on both carbon reserves and current photoassimilates. The different responses of the two Quercus species to temperature and photoperiod related more to xylem structure than to the source of carbon used for budburst.     

Fossils of the Fagaceae and their implications in systematics and biogeography

The fossil history of the Fagaceae from China and its systematic and biogeographic implications are discussed based on revisionary studies of the fossil records. No creditable macrofossil record of the Fagaceae exists in the Cretaceous deposits and all the Cretaceous microfossil reports remain equivocal and require further study. The Paleocene fossils show the appearance and diversification of the two groups corresponding to the subfamilies Fagoideae and Castaneoideae sensu Nixon. By the Eocene, all modern genera had been present. The oldest fagaceous fossils represent subfamily Fagoideae with affinities to the extant genus Trigonobalanus. The leaf fossil genus Berryophyllum, with affinities to Quercus subg. Cyclobalanopsis, has been documented by the early Eocene and might have occurred earlier than other fossils assignable to Quercus. The appearance of evergreen sclerophyllous Ouercus with entire leaves might have occurred earlier than those with toothed leaves. Deciduous, urticoid-leaved oak fossils (Quercus subg. Quercus sect. Quercus) had not appeared until the Miocene. Fossil equivalents of Trigonobalanus, Castanopsis and Lithocarpus had occurred in Europe and North America by the early Tertiary, suggesting that continuous distributions were achieved via the northern hemisphere land bridges. Three groups of evergreen sclerophyllous oaks of apparent close phylogenetic relationships occurred in the Hengduan mountains, the Mediterranean area and northwestern North America. Their fossil forms have become dominant elements of those vegetation zones since the Miocene. A shared fossil history indicates a possible biogeographic boundary formed by the ancient Mediterranean. The evidence suggests that the oaks might arrive in North America during two distinct geologic periods: evergreen sclerophyllous entire-leaved oaks appeared by the Early Tertiary, whereas thedeciduous oaks with urticoid leaves appeared in the Late Tertiary.     

壳斗科的地质历史及其系统学和植物地理学意义

在收集整理现有壳斗科化石资料的基础上,讨论了壳斗科及其各属的起源时间、地史分布和地史 演替过程以及这些化石资料在系统学和植物地理学上的意义。白垩纪尚无壳斗科可靠的大化石记录, 微化石需要进一步研究才能确定亲缘关系以及古新世壳斗科已经分化出两个类群。从以上这些事实推 论壳斗科起源于白垩纪晚期,而壳斗科现代各属出现的时间应不晚于古新世。最早发现的壳斗科化石和现代栗亚科和水青冈亚科在形态结构上非常相似,这一事实表明,壳斗科分为两个亚科的观点更接近客观事实。在水青冈亚科中,三棱栎类的化石最早出现;在栎属中,青冈亚属更接近祖先类群;在地史中全缘栎类较具齿栎类出现早,粗齿的落叶栎类出现最晚。三棱栎属、栲属和石栎属的化石在老第三纪出现于北美和欧洲的事实说明,北美、欧洲和东亚在老第三纪时有一个相通的壳斗科植物区系。南美的三棱栎是通过北美进入南美的。中国横断山、欧洲地中海沿岸和北美西北部有一类形态特征相似、亲缘关系相近的硬叶栎类,它们之间有相同的地质演替历史,它们现代分布边界可能就是古地中海的边界。美洲的栎类有两个来源,常绿硬叶栎类是通过古地中海沿岸而经北美-欧洲陆桥到达的,落叶栎类则是在中新世以后通过白令海峡到达的。     

Stem xylem features in three Quercus (Fagaceae) species along a climatic gradient in NE Spain

 Stem xylem features in two evergreen Quercus species (Q. coccifera and Q. ilex) and a deciduous one (Q. faginea) were analysed along an Atlantic-Mediterranean climatic gradient in which rainfall and winter cold experience strong variation. Mean maximum vessel diameter, vessel density, vessel element length, xylem transverse sectional area, Huber value (xylem transverse sectional area per leaf area unit), theoretical leaf specific conductivity (estimated hydraulic conductance per leaf area unit) and total leaf area were determined in 3-year-old branches. Q. faginea presented the widest vessels and the highest theoretical leaf specific conductivity while Q. coccifera showed the lowest total leaf area and the highest Huber value. Studied features did not exhibit significant correlations with mean minimum January temperature in any species but did show significant relationships with rainfall. In Q. coccifera, mean maximum vessel diameter, vessel element length and theoretical leaf specific conductivity increased with higher rainfall while vessel density decreased. Mean maximum vessel diameter and total leaf area in Q. ilex increased with precipitation whereas variables of Q. faginea did not show any significant trend. Results suggest that aridity, rather than minimum winter temperature, controls stem xylem responses in the studied evergreen species. Q. faginea traits did not show any response to precipitation, probably because this species develops deep roots, which in turn makes edaphic and topographic factors more important in the control of soil water availability. In response to aridity Q. coccifera only exhibits adjustment at a xylem level by reducing its water transport capacity through a reduction of vessel diameter without changing the amount of xylem tissue or foliage, whereas Q. ilex adjusts its water transport capacity in parallel to the foliage area. Received: 13 January 1997 / Accepted: 8 April 1997     

Mechanism of freeze-induced embolism in Fagus sylvatica L.

 The mechanism of freeze stress-induced embolism in Fagus sylvatica L. branches was analyzed under controlled conditions. Excised branches were exposed to successive freeze-thaw cycles in temperature controlled chambers. Thermocouples were placed on the bark to detect sap freezing exotherms. The degree of xylem embolism was estimated after each cycle by the loss of hydraulic conductivity. After one freeze-thaw cycle the degree of embolism was found to decrease with xylem specific hydraulic conductivity, small apical shoots being more susceptible to embolism. Exotherms revealed that distal shoots were freezing first and exuded sap as a result of water expansion. The lower water content in apical shoots upon freezing probably induced higher sap tensions which promoted air bubble expansion and vessel cavitation preferentially near the apices. When the decrease in water content was experimentally prevented, embolism developed to a lesser extent. The higher vulnerability of shoot apices may protect the rest of the branch from winter damage. Received: 29 May 1998 / Accepted: 15 August 1998     

Aboveground biomass allocation patterns within Mediterranean sub-shrubs: A quantitative analysis of seasonal dimorphism

The monthly patterns of aboveground biomass allocation were studied in the branches of six Mediterranean sub-shrubs with different leaf phenology. Four of them were seasonally dimorphic species, and the remaining two were a winter deciduous and a cushion plant with photosynthetic stems. By the analysis of these species we aimed to identify different aboveground biomass allocation patterns within seasonally dimorphic species and to understand the role of seasonal dimorphism as a strategy to avoid the main stresses of mediterranean climate: summer drought and winter cold. The biomass allocation to the different living and photosynthetic fractions of 3-year-old branches was studied monthly for a minimum of 13 months per species. Leaf area (LA, mm²) and leaf mass per area (LMA, mg cm⁻²) measurements were used to characterize the diverse types of leaves of each species. Standing dead and senescent tissues accounted for a great percentage of the branch biomass of seasonally dimorphic species both during summer and winter. Different patterns of photosynthetic biomass allocation were found within the seasonally dimorphic species analysed. These patterns ranged from the moderate photosynthetic biomass oscillation of Salvia lavandulifolia to the almost deciduousness of Lepidium subulatum, and they were achieved by keeping alive, drying out or shedding different types of branches and leaves throughout the year. The formation of stress tolerant leaves and the reduction in the amount of photosynthetic biomass responded both to the occurrence of summer drought and winter cold. These results demonstrate that seasonal dimorphism is a flexible ecological strategy, as it comprises very different leaf phenologies and enables plants to escape both summer drought and winter cold.     

Evidence for xylem embolism as a primary factor in dehydration-induced declines in leaf hydraulic conductance

Hydraulic conductance of leaves (K(leaf)) typically decreases with increasing water stress and recent studies have proposed different mechanisms responsible for decreasing K(leaf) . We measured K(leaf) concurrently with ultrasonic acoustic emissions (UAEs) in dehydrating leaves of several species to determine whether declining K(leaf) was associated with xylem embolism. In addition, we performed experiments in which the surface tension of water in the leaf xylem was reduced by using a surfactant solution. Finally, we compared the hydraulic vulnerability of entire leaves with the leaf lamina in three species. Leaf hydraulic vulnerability based on rehydration kinetics and UAE was very similar, except in Quercus garryana. However, water potentials corresponding to the initial decline in K(leaf) and the onset of UAE in Q. garryana were similar. In all species tested, reducing the surface tension of water caused K(leaf) to decline at less negative water potentials compared with leaves supplied with water. Microscopy revealed that as the fraction of embolized xylem increased, K(leaf) declined sharply in Q. garryana. Measurements on leaf discs revealed that reductions in lamina hydraulic conductance with dehydration were not as great as those observed in intact leaves, suggesting that embolism was the primary mechanism for reductions in K(leaf) during dehydration.     

Seasonal photosynthetic changes in the green-stemmed Mediterranean shrub <Emphasis Type="Italic">Calicotome villosa</Emphasis>: a comparison with leaves

Some photosynthetic attributes of leaves and stems were seasonally followed in the small-leaved, summer-deciduous, green-stemmed Mediterranean shrub Calicotome villosa. Both leaves and stems displayed similar photon energy-saturated photosystem 2 (PS2) efficiencies with a minimum during winter. A second minimum in stems during the leafless summer period could be ascribed to sustained photoinhibition. Yet, stems were slightly inferior in photon capture, resulting partly from lower chlorophyll (Chl) contents and partly from higher reflectance due to pubescence. As a result, photon energy-saturated linear electron transport rates were slightly higher in leaves. However, when the total leaf and stem areas were taken into account, this superiority was abolished during autumn and winter and more than overturned during spring. Given that during summer the stems were the only photosynthetic organs, the yearly photosynthetic contribution of stems was much higher. Chl contents in stems displayed a transient and considerable summer drop, accompanied by an increase in the carotenoid to Chl ratio, indicating a photo-protective adaptation to summer drought through a decrease of photo-selective capacity, typical for leaves of many Mediterranean plants.     

Hydraulic properties and freezing-induced cavitation in sympatric evergreen and deciduous oaks with contrasting habitats

We investigated the hydraulic properties in relation to soil moisture, leaf habit, and phylogenetic lineage of 17 species of oaks (Quercus) that occur sympatrically in northern central Florida (USA). Leaf area per shoot increased and Huber values (ratio of sapwood area to leaf area) decreased with increasing soil moisture of species’ habitats. As a result, maximum hydraulic conductance and maximum transpiration were positively correlated with mean soil moisture when calculated on a sapwood area basis, but not when calculated on a leaf area basis. This reveals the important role that changes in allometry among closely related species can play in co‐ordinating water transport capacity with soil water availability. There were significant differences in specific conductivity between species, but these differences were not explained by leaf habit or by evolutionary lineage. However, white oaks had significantly smaller average vessel diameters than red oaks or live oaks. Due to their lower Huber values, maximum leaf specific conductivity (K_L) was higher in evergreen species than in deciduous species and higher in live oaks than in red oaks or white oaks. There were large differences between species and between evolutionary lineages in freeze–thaw‐induced embolism. Deciduous species, on average, showed greater vulnerability to freezing than evergreen species. This result is strongly influenced by evolutionary lineage. Specifically, white oaks, which are all deciduous, had significantly higher vulnerability to freezing than live oaks (all evergreen) and red oaks, which include both evergreen and deciduous species. These results highlight the importance of taking evolutionary lineage into account in comparative physiological studies.     

Hydraulic disruption and passive migration by a bacterial pathogen in oak tree xylem

Xylella fastidiosa (Xf) is a xylem-limited bacterial pathogen that causes leaf scorch symptoms in numerous plant species in urban, agricultural, and natural ecosystems worldwide. The exact mechanism of hydraulic disruption and systemic colonization of xylem by Xf remains elusive across all host plants. To understand both processes better, the functional and structural characteristics of xylem in different organs of both healthy and Xf-infected trees of several Quercus species were studied. Hydraulic conductivity (K(s)) in Xf-infected petioles of Q. palustris and Q. rubra decreased significantly compared with healthy trees as the season progressed and plummeted to zero with the onset of scorch symptoms. Prior to the onset of symptoms, embolism was as much as 3.7 times higher in Xf-infected petioles compared with healthy controls and preceded significant reductions in K(s). Embolism likely resulted from pit membrane degradation during colonization of new petiole xylem and triggered the process that eventually led to vessel occlusion. Pit membrane porosity was studied using the following four methods to determine if a pathway exists in the xylem network of woody stems that allows for passive Xf migration: (i) calculations based on vulnerability to cavitation data, (ii) scanning electron micrographs, (iii) microsphere injections, and (iv) air seeding thresholds on individual vessels. All four methods consistently demonstrated that large pit membrane pores (i.e. greater than the diameter of individual Xf) occur frequently throughout the secondary stem xylem in several Quercus species. These large pores probably facilitate systemic colonization of the secondary xylem network and contribute to the high susceptibility to bacterial leaf scorch exhibited among these species.     

Functional lifespans of xylem vessels: Development,hydraulic function,and post‐function of vessels in several species of woody plants

Premise of the Study

Xylem vessels transition through different stages during their functional lifespan, including expansion and development of vessel elements, transition to vessel hydraulic functionality, and eventual transition to post‐functionality. We used information on vessel development and function to develop a model of vessel lifespan for woody plants.

Methods

We examined vessel functional lifespan using repeated anatomical sampling throughout the growing season, combined with active‐xylem staining to evaluate vessel hydraulic transport functionality. These data were combined with a literature review. The transitions between vessel functional lifespans for several species are illustrated, including grapevine (Vitis vinifera L., Vitaceae), English oak (Quercus robur L., Fagaceae), American chestnut [Castanea dentata (Marshall) Borkh.; Fagaceae], and several arid and semi‐arid shrub species.

Key Results

In intact woody plants, development and maturation of vessel elements may be gradual. Once hydraulically functional, vessel elements connect to form a vessel network that is responsible for bulk hydraulic flow through the xylem. Vessels become nonfunctional due to the formation of gas emboli. In some species and under some conditions, vessel functionality of embolized conduits may be restored through refilling. Blockages, such as tyloses, gels, or gums, indicate permanent losses in hydraulic functional capacity; however, there may be some interesting exceptions to permanent loss of functionality for gel‐based blockages.

Conclusions

The gradual development and maturation of vessel elements in woody plants, variation in the onset of functionality between different populations of vessels throughout the growing season, and differences in the timing of vessel transitions to post‐functionality are important aspects of plant hydraulic function.     

Rapid hydraulic recovery in Eucalyptus pauciflora after drought: linkages between stem hydraulics and leaf gas exchange

In woody plants, photosynthetic capacity is closely linked to rates at which the plant hydraulic system can supply water to the leaf surface. Drought‐induced embolism can cause sharp declines in xylem hydraulic conductivity that coincide with stomatal closure and reduced photosynthesis. Recovery of photosynthetic capacity after drought is dependent on restored xylem function, although few data exist to elucidate this coordination. We examined the dynamics of leaf gas exchange and xylem function in Eucalyptus pauciflora seedlings exposed to a cycle of severe water stress and recovery after re‐watering. Stomatal closure and leaf turgor loss occurred at water potentials that delayed the extensive spread of embolism through the stem xylem. Stem hydraulic conductance recovered to control levels within 6 h after re‐watering despite a severe drought treatment, suggesting an active mechanism embolism repair. However, stomatal conductance did not recover after 10 d of re‐watering, effecting tighter control of transpiration post drought. The dynamics of recovery suggest that a combination of hydraulic and non‐hydraulic factors influenced stomatal behaviour post drought.     

Morphological and physiological modifications of Cistus salvifolius L. winter leaves in response to the rise in winter temperatures

The global climate is predicted to change in the next century; for the Mediterranean Basin, an increase in air temperature more than 4°C and a higher frequency of extreme climatic events such as drought and heat waves are expected. In this work, the response of Cistus salvifolius L. to the rise in winter temperature has been studied. Plants acclimated to winter conditions [outdoor (OUT)] were moved into a greenhouse [indoor (IND)] at higher temperature and eco-physiological behaviour was analysed on leaves after 15 days from plant transferring (IND_15d) and on leaves developed IND. IND leaves were characterized by reduced thickness, higher specific leaf area, higher CO₂ mesophyll conductance and photosynthetic rate, and lower respiratory rate than leaves grown OUT upon current winter conditions. In IND_15d leaves, no improvement of photochemical activity was found. When IND leaves were subjected to a rapid increase in air temperature, the CO₂ fixation was not limited indicating a high thermotolerance of photosynthetic machinery. The results for IND leaves indicate the occurrence of a strategy that merging changes in leaf structure as well as in photosynthetic regulation allow C. salvifolius to maintain an elevated carbon gain in response to temperature increase.     

Sixteen years of winter stress: an assessment of cold hardiness,growth performance and survival of hybrid poplar clones at a boreal planting site

In recent years, thousands of hectares of hybrid poplar plantations have been established in Canada for the purpose of carbon sequestration and wood production. However, boreal planting environments pose special challenges that may compromise the long‐term survival and productivity of such plantations. In this study, we evaluated the effect of winter stress, that is, frequent freeze‐thaw and extreme cold events, on growth and survival of 47 hybrid poplar clones in a long‐term field experiment. We further assessed physiological and structural traits that are potentially important for cold tolerance for a selected set of seven clones. We found that trees with narrow xylem vessels showed reduced freezing‐induced embolism and showed superior productivity after 16 growing seasons. With respect to cold hardiness of living tissues, we only observed small differences among clones in early autumn, which were nonetheless significantly correlated to growth. Maximum winter cold hardiness and the timing of leaf senescence and budbreak were not related to growth or survival. In conclusion, our data suggest that reduction of freezing‐induced embolism due to small vessel diameters is an essential adaptive trait to ensure long‐term productivity of hybrid poplar plantations in boreal planting environments.