Does freezing and dynamic flexing of frozen branches impact the cavitation resistance of Malus domestica and the Populus clone Walker?

Frost damage to the xylem conduits of trees is a phenomenon of eco-physiological importance. It is often documented in terms of the percentage loss of conductivity (PLC), an indicator of air filling of the conduits. However, trees that refill their conduits in spring could be impacted more by damage to the conduits that reduce cavitation resistance, making them more susceptible to future drought events. We investigated whether ice formation, dynamic flexing of frozen branches or freeze–thaw events could reduce the cavitation resistance (cause “frost fatigue”) in first-year shoots of apple (Malus domestica) and clonal hybrid cottonwood (Walker). Frost fatigue was measured in terms of P₅₀ (the negative xylem pressure required to cause a 50 % loss of conductivity). All treatment groups showed significant frost fatigue, with the exception of the pre-flushed, constantly frozen poplar branches. The P₅₀ following freeze treatments was approximately 50 % of the pre-freeze values. The effect tended to be greater in freeze–thawed branches. Dynamic bending of the branches had no effect on either PLC or P₅₀. In three out of four cases, there was a significant correlation between P₅₀ and PLC. Frost fatigue occurred in both apple and poplar, two unrelated species with different drought and frost tolerances, suggesting that it may be a widespread phenomenon that could impact the ecophysiology of temperate forests.     

Poplar vulnerability to xylem cavitation acclimates to drier soil conditions

Xylem vulnerability to cavitation differs between tree species according to their drought resistance, more xerophilous species being more resistant to xylem cavitation. Variability in xylem vulnerability to cavitation is also found within species, especially between in situ populations. The origin of this variability has not been clearly identified. Here we analyzed the response of xylem hydraulic traits of Populus tremula×Populus alba trees to three different soil water regimes. Stem xylem vulnerability was scored as the xylem water potential causing 12, 50 and 88% loss of conductivity (P₁₂, P₅₀ and P₈₈). Vulnerability to cavitation was found to acclimate to growing conditions under different levels of soil water content, with P₅₀ values of ?1.82, ?2.03 and ?2.45 MPa in well‐watered, moderately water‐stressed and severely water‐stressed poplars, respectively. The value of P₁₂, the xylem tension at which cavitation begins, was correlated with the lowest value of midday leaf water potential (ψm) experienced by each plant, the difference between the two parameters being approximately 0.5 MPa, consistent with the absence of any difference in embolism level between the different water treatments. These results support the hypothesis that vulnerability to cavitation is a critical trait for resistance to drought. The decrease in vulnerability to cavitation under growing conditions of soil drought was correlated with decreased vessel diameter, increased vessel wall thickness and a stronger bordered pit field (t/b)². The links between these parameters are discussed.     

Vulnerability to drought‐induced cavitation in poplars: synthesis and future opportunities

Vulnerability to drought‐induced cavitation is a key trait of plant water relations. Here, we summarize the available literature on vulnerability to drought‐induced cavitation in poplars (Populus spp.), a genus of agronomic, ecological and scientific importance. Vulnerability curves and vulnerability parameters (including the water potential inducing 50% loss in hydraulic conductivity, P₅₀) were collected from 37 studies published between 1991 and 2014, covering a range of 10 species and 12 interspecific hybrid crosses. Results of our meta‐analysis confirm that poplars are among the most vulnerable woody species to drought‐induced cavitation (mean P₅₀ = ?1.44 and ?1.55 MPa across pure species and hybrids, respectively). Yet, significant variation occurs among species (P₅₀ range: 1.43 MPa) and among hybrid crosses (P₅₀ range: 1.12 MPa), within species and hybrid crosses (max. P₅₀ range reported: 0.8 MPa) as well as in response to environmental factors including nitrogen fertilization, irradiance, temperature and drought (max. P₅₀ range reported: 0.75 MPa). Potential implications and gaps in knowledge are discussed in the context of poplar cultivation, species adaptation and climate modifications. We suggest that poplars represent a valuable model for studies on drought‐induced cavitation, especially to elucidate the genetic and molecular basis of cavitation resistance in Angiosperms.     

Seasonality of cavitation and frost fatigue in Acer mono Maxim.

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

The physiological implications of primary xylem organization in two ferns

Xylem structure and function are well described in woody plants, but the implications of xylem organization in less‐derived plants such as ferns are poorly understood. Here, two ferns with contrasting phenology and xylem organization were selected to investigate how xylem dysfunction affects hydraulic conductivity and stomatal conductance (g_s). The drought‐deciduous pioneer species, Pteridium aquilinum, exhibits fronds composed of 25 to 37 highly integrated vascular bundles with many connections, high g_s and moderate cavitation resistance (P₅₀ = ?2.23 MPa). By contrast, the evergreen Woodwardia fimbriata exhibits sectored fronds with 3 to 5 vascular bundles and infrequent connections, low g_s and high resistance to cavitation (P₅₀ = ?5.21 MPa). Xylem‐specific conductivity was significantly higher in P. aqulinium in part due to its wide, efficient conduits that supply its rapidly transpiring pinnae. These trade‐offs imply that the contrasting xylem organization of these ferns mirrors their divergent life history strategies. Greater hydraulic connectivity and g_s promote rapid seasonal growth, but come with the risk of increased vulnerability to cavitation in P. aquilinum, while the conservative xylem organization of W. fimbriata leads to slower growth but greater drought tolerance and frond longevity.     

Water loss regulation to soil drought associated with xylem vulnerability to cavitation in temperate ring-porous and diffuse-porous tree seedlings

Key message

Sustainable stomatal opening despite xylem cavitation occurs in ring-porous species and stomatal closure prior to cavitation in diffuse-porous species during soil drought.

Abstract

To elucidate the relationship between water loss regulation and vulnerability to cavitation associated with xylem structure, stomatal conductance (g _s), defoliation, vulnerability curves, and vessel features were measured on seedlings of ring-porous Zelkova serrata and Melia azedarach, and diffuse-porous Betula platyphylla var. japonica, Cerasus jamasakura and Carpinus tschonoskii. Under prolonged drought conditions, the percentage loss of hydraulic conductivity (PLC) increased and g _s decreased gradually with decreasing predawn (Ψ_pd) or xylem water potential (Ψ_xylem) in Z. serrata. During the gentle increase of PLC in M. azedarach, g _s increased in the early stages of dehydration while leaves were partly shed. A sharp reduction in g _s was observed before the onset of an increase in the PLC for drying plants of the three diffuse-porous species, suggesting cavitation avoidance by stomatal regulation. In the ring-porous species, xylem-specific hydraulic conductivity (K _s) was higher, whereas the vessel multiple fractions, the ratio of the number of grouped vessels to total vessels, was lower than that in the diffuse-porous species, suggesting that many were distributed as solitary vessels. This may explain the gradual increase in the PLC with decreasing Ψ_xylem because isolated vessels provide less opportunity for air seeding. Different water loss regulation to soil drought was identified among the species, with potential mechanisms being sustainable gas exchange at the expense of xylem dysfunction or partial leaf shedding, and the avoidance of xylem cavitation by strict stomatal regulation. These were linked to vulnerability to cavitation that appears to be governed by xylem structural properties.     

Water relations of Robinia pseudoacacia L.: do vessels cavitate and refill diurnally or are R‐shaped curves invalid in Robinia?

Since 2005, an unresolved debate has questioned whether R‐shaped vulnerability curves (VCs) might be an artefact of the centrifuge method of measuring VCs. VCs with R‐shape show loss of stem conductivity from approximately zero tension, and if true, this suggests that some plants either refill embolized vessels every night or function well with a high percentage of vessels permanently embolized. The R‐shaped curves occur more in species with vessels greater than half the length of the segments spun in a centrifuge. Many have hypothesized that the embolism is seeded by agents (bubbles or particles) entering the stem end and travelling towards the axis of rotation in long vessels, causing premature cavitation. VCs were measured on Robinia pseudoacacia L. by three different techniques to yield three different VCs; R‐shaped: Cavitron P₅₀ = 0.30 MPa and S‐shaped: air injection P₅₀ = 1.48 MPa and bench top dehydration P₅₀ = 3.57 MPa. Stem conductivity measured in the Cavitron was unstable and is a function of vessel length when measured repeatedly with constant tension, and this observation is discussed in terms of stability of air bubbles drawn into cut‐open vessels during repeated Cavitron measurement of conductivity; hence, R‐shaped curves measured in a Cavitron are probably invalid.     

Hydraulic vulnerability, vessel refilling, and seasonal courses of stem water potential of Sorbus aucuparia L. and Sambucus nigra L.

Differences in the seasonal variation in stem water potential between the two shrub species Sorbus aucuparia and Sambucus nigra were related with their vulnerability to xylem cavitation. It was also demonstrated indirectly that the two species differ in the extent to which they reverse cavitation. Seasonal variation in stem water potential was investigated during three growing seasons with in situ stem psychrometers. Sorbus experienced wide water potential variations and reached a minimum of -4.2 MPa during drought. Under the same microclimatic conditions, Sambucus experienced consistent stem water potentials with a minimum of -1.7 MPa. The relationship between percentage loss in hydraulic conductivity (PLC) and water potential (hydraulic vulnerability curve) of the two species differed in shape: a flat curve with nearly total loss of conductivity at -6 MPa was found for SORBUS: Sambucus showed a steep vulnerability curve with 90% loss conductivity at -2.2 MPa. Thus, Sambucus is extremely vulnerable to cavitation, but Sorbus is an almost invulnerable species. This different cavitation resistance adjusted the ranges of field stem water potential that the species experienced. Finally, seasonal courses of naturally occurring (native) embolism were compared with calculated PLC courses. This comparison indicates that Sorbus did not refill embolized xylem vessels whereas Sambucus reversed embolism. It was concluded that species which are highly vulnerable to cavitation and drought-induced embolism need refilling of embolized vessels as well as isohydric water potential patterns as two strategies of survival.     

Vulnerability to xylem cavitation of Hakea species (Proteaceae) from a range of biomes and life histories predicted by climatic niche

Background and AimsExtreme drought conditions across the globe are impacting biodiversity, with serious implications for the persistence of native species. However, quantitative data on physiological tolerance are not available for diverse flora to inform conservation management. We quantified physiological resistance to cavitation in the diverse Hakea genus (Proteaceae) to test predictions based on climatic origin, life history and functional traits.MethodsWe sampled terminal branches of replicate plants of 16 species in a common garden. Xylem cavitation was induced in branches under varying water potentials (tension) in a centrifuge, and the tension generating 50 % loss of conductivity (stem P₅₀) was characterized as a metric for cavitation resistance. The same branches were used to estimate plant functional traits, including wood density, specific leaf area and Huber value (sap flow area to leaf area ratio).Key ResultsThere was significant variation in stem P₅₀ among species, which was negatively associated with the species climate origin (rainfall and aridity). Cavitation resistance did not differ among life histories; however, a drought avoidance strategy with terete leaf form and greater Huber value may be important for species to colonize and persist in the arid biome.ConclusionsThis study highlights climate (rainfall and aridity), rather than life history and functional traits, as the key predictor of variation in cavitation resistance (stem P₅₀). Rainfall for species origin was the best predictor of cavitation resistance, explaining variation in stem P₅₀, which appears to be a major determinant of species distribution. This study also indicates that stem P₅₀ is an adaptive trait, genetically determined, and hence reliable and robust for predicting species vulnerability to climate change. Our findings will contribute to future prediction of species vulnerability to drought and adaptive management under climate change.     

Ecological differentiation in xylem cavitation resistance is associated with stem and leaf structural traits

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P₅₀) related to dry season leaf water potentials and stem and leaf traits. P₅₀‐values ranged from ?0.8 to ?6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade‐tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber‐value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade‐off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P₅₀, but pioneers and deciduous species had smaller hydraulic safety margins than shade‐tolerants and evergreens. A trade‐off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species.     

Measurement of vulnerability to water stress‐induced cavitation in grapevine: a comparison of four techniques applied to a long‐vesseled species

Among woody plants, grapevines are often described as highly vulnerable to water‐stress induced cavitation with emboli forming at slight tensions. However, we found native embolism never exceeded 30% despite low xylem water potentials (Ψ_x) for stems of field grown vines. The discrepancy between native embolism measurements and those of previous reports led us to assess vulnerability curve generation using four separate methods and alterations (i.e. segment length and with/without flushing to remove embolism prior to measurement) of each. Centrifuge, dehydration and air‐injection methods, which rely on measurement of percentage loss of hydraulic conductivity (PLC) in detached stems, were compared against non‐invasive monitoring of xylem cavitation with nuclear magnetic resonance (NMR) imaging. Short segment air‐injection and flushed centrifuge stems reached >90 PLC at Ψ_x of‐0.5 and ?1.5 MPa, respectively, whereas dehydration and long‐segment air‐injection measurements indicated no significant embolism at Ψ_x > ?2.0 MPa. Observations from NMR agreed with the dehydration and long segment air‐injection methods, showing the majority of vessels were still water‐filled at Ψ_x > ?1.5 MPa. Our findings show V. vinifera stems are far less vulnerable to water stress‐induced cavitation than previously reported, and dehydration and long segment air‐injection techniques are more appropriate for long‐vesseled species and organs.     

Hydraulic conductance and vulnerability to cavitation in corn (Zea mays L.) hybrids of differing drought resistance

Water transport through the xylem is essential for replacing water loss during transpiration, thus preventing desiccation and permitting photosynthesis. The occurrence of cavitation and embolism due to drought impairs transport to the transpiring leaves. Most research in this discipline has been conducted on woody plants. Less attention has been given to cavitation of crops and its physiological significance for understanding crop water relations. In this paper, hydraulic conductance and vulnerability of xylem to cavitation were studied in corn hybrids with different drought resistances. The results indicated that stems of drought-resistant ‘Pioneer 3902’ not only had a higher conductivity on both a stem area and leaf area basis but also had a greater resistance to cavitation. The estimated xylem pressure at 63.2% loss of conductivity (Weibull fitting parameter b) and at 50% loss of conductivity (P₅₀) in ‘Pioneer 3902’ were about 0.2 MPa lower than in ‘Pride 5’. Higher conductivity in ‘Pioneer 3902’ was mainly attributed to more vascular bundles per stem area rather than greater vessel diameter. The central bundles and peripheral vascular bundles showed the same degree of cavitation although the vessels of central bundles were generally larger than in peripheral bundles.     

Excising stem samples underwater at native tension does not induce xylem cavitation

Xylem resistance to water stress‐induced cavitation is an important trait that is associated with drought tolerance of plants. The level of xylem cavitation experienced by a plant is often assessed as the percentage loss in conductivity (PLC) at different water potentials. Such measurements are constructed with samples that are excised underwater at native tensions. However, a recent study concluded that cutting conduits under significant tension induced cavitation, even when samples were held underwater during cutting. This resulted in artificially increased PLC because of what we have termed a ‘tension‐cutting artefact’. We tested the hypothesized tension‐cutting artefact on five species by measuring PLC at native tension compared with after xylem tensions had been relaxed. Our results did not support the tension‐cutting artefact hypothesis, as no differences were observed between native and relaxed samples in four of five species. In a fifth species (Laurus nobilis), differences between native and relaxed samples appear to be due to vessel refilling rather than a tension‐cutting effect. We avoided the tension‐cutting artefact by cutting samples to slightly longer than their measurement length and subsequent trimming of at least 0.5 cm of sample ends prior to measurement.     

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

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

Prediction of hydraulic conductivity loss from relative water loss: new insights into water storage of tree stems and branches

More frequently occurring, drought waves call for a deeper understanding of tree hydraulics and fast and easily applicable methods to measure drought stress. The aim of this study was to establish empirical relationships between the percent loss of hydraulic conductivity (PLC) and the relative water loss (RWL) in woody stem axes with different P₅₀, i.e. the water potential (Ψ) that causes 50% conductivity loss. Branches and saplings of temperate conifer (Picea abies, Larix decidua) and angiosperm species (Acer campestre, Fagus sylvatica, Populus x canescens, Populus tremula, Sorbus torminalis) and trunk wood of mature P. abies trees were analyzed. P₅₀ was calculated from hydraulic measurements following bench top dehydration or air injection. RWL and PLC were fitted by linear, quadratic or cubic equations. Species‐ or age‐specific RWLs at P₅₀ varied between 10 and 25% and P₈₈, the Ψ that causes 88% conductivity loss, between 18 and 44%. P₅₀ was predicted from the relationship between Ψ and the RWL. The predictive quality for P₅₀ across species was almost 1:1 (r² = 0.99). The approach presented allows thus reliable and fast prediction of PLC from RWL. Branches and saplings with high hydraulic vulnerability tended to have lower RWLs at P₅₀ and at P₈₈. The results are discussed with regard to the different water storage capacities in sapwood and survival strategies under drought stress. Potential applications are screening trees for drought sensitivity and a fast interpretation of diurnal, seasonal or drought induced changes in xylem water content upon their impact on conductivity loss.     

Cavitation fatigue in conifers: a study on eight European species

After drought-induced embolism and repair, tree xylem may be weakened against future drought events (cavitation fatigue). As there are few data on cavitation fatigue in conifers available, we quantified vulnerability curves (VCs) after embolism/repair cycles on eight European conifer species. We induced 50% and 100% loss of conductivity (LC) with a cavitron, and analyzed VCs. Embolism repair was obtained by vacuum infiltration. All species demonstrated complete embolism repair and a lack of any cavitation fatigue after 50% LC . After 100% LC, European larch (Larix decidua), stone pine (Pinus cembra), Norway spruce (Picea abies), and silver fir (Abies alba) remained unaffected, while mountain pine (Pinus mugo), yew (Taxus baccata), and common juniper (Juniperus communis) exhibited 0.4–0.9 MPa higher vulnerability to embolism. A small cavitation fatigue observed in Scots pine (Pinus sylvestris) was probably biased by incomplete embolism repair, as indicated by a correlation of vulnerability shifts and conductivity restoration. Our data demonstrate that cavitation fatigue in conifers is species-specific and depends on the intensity of preceding LC. The lack of fatigue effects after moderate LC, and relevant effects in only three species after high LC, indicate that conifers are relatively resistant against cavitation fatigue. This is remarkable considering the complex and delicate conifer pit architecture and may be important considering climate change projections.     

Water stress induced cavitation and embolism in some woody plants

A comparison was made of the relative vulnerability of xylem conduits to cavitation and embolism in three species [ Thuja occidentalis L., Tsuga canadensis (L.) Carr. and Acer saccharum Marsh.]. Waterlogged samples of wood were air dehydrated while measuring relative water loss, loss of hydraulic conductance, cumulative acoustic emissions (= cavitations) and xylem water potential. Most cavitation events and loss of hydraulic conductance occurred while water potential declined from – 1 to –6 MPa. There were differences in vulnerability between species. Other people have hypothesized that large xylem conduits (e.g. vessels) should be more vulnerable to cavitations than small conduits (e.g. tracheids). Our findings are contrary to this hypothesis. Under water stress, the vessel bearing wood retained water better than tracheid bearing wood. However, within a species large conduits were more prone to cavitation than small conduits.     

Phenotypic plasticity in mesic populations of Pinus pinaster improves resistance to xylem embolism (P50) under severe drought

The objectives of the study were to assess the phenotypic variation in the vulnerability to water stress-induced cavitation (estimated by P₅₀, or the xylem water potential which causes a 50% loss of conductivity) and the trade-offs between P₅₀ and related hydraulic traits, i.e., stem specific conductivity (K _s), slope of the vulnerability curve (slope), wood density and branch size. Variability was examined for six Pinus pinaster populations covering the latitudinal range of the species and plasticity was tested through two provenance-progeny trial sites (xeric/mesic). As expected, the overall values of P₅₀, K _s and branch size decreased in the xeric site. Variation in P₅₀ and K _s among populations was mainly the result of phenotypic plasticity, while wood density was genetically controlled and not affected by the environment. Stress conditions in the xeric site promoted a convergence in P₅₀ and K _s as a result of the high phenotypic plasticity of the populations from mesic origins. In the mesic site, the ranking of populations for cavitation resistance and hydraulic capacity was consistent with the geographic location of the seed source. Higher resistance to cavitation was related to lower K _s, branch size and slope, mainly at the population level, but also as a general trend across individuals. In a warmer and drier climate, there could be a potential selection of Pinus pinaster populations from mesic origins, which showed a great responsiveness and adjustment to drought conditions (similar or higher P₅₀ than the populations from dry origins), in addition to a high wood density and growth.     

Cavitation fatigue and its reversal in sunflower (Helianthus annuus L.)

"Cavitation fatigue" is the increased susceptibility of a xylem conduit to cavitation as a result of its prior cavitation. It was investigated whether cavitation fatigue induced in vivo could be repaired in intact plants. Sunflowers (Helianthus annuus L.) were subjected to soil drought in the greenhouse. Native embolism and vulnerability to cavitation was measured in well-watered controls and after 5 d and 10 d of controlled drought. A dramatic cavitation fatigue was observed where droughted xylem that was refilled in the laboratory developed up to 60 PLC (percentage loss of hydraulic conductivity) at -1 MPa versus only 5.2 PLC in non-droughted controls. Rewatered plants showed the complete reversal of cavitation fatigue over 4 d. Reversal of fatigue was correlated with the refilling of embolized vessels in the intact plants (r(2)=0.91, P<0.01), suggesting that xylem transport to fatigued vessels was required for their repair. The in vivo reversal of fatigue was partially duplicated in excised stem segments by perfusing them with root exudates from droughted (DR) and well-watered (WW) plants. The DR exudate had a greater effect, and this was associated with a greater pH in the DR versus WW saps, but there was no difference in total cation concentration. Perfusions with 2 mM CaCl(2) and KCl solutions also partially reversed cavitation fatigue as opposed to no effect with deionized water, suggesting a role of ions in addition to a pH effect. It is suspected that fatigue is caused by stretching and partial disruption of linkages between cellulose microfibrils in inter-conduit pit membranes during air seeding, and that the reversal of fatigue involves restoring these linkages by ingredients in xylem sap.     

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

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