Chaparral is the most abundant vegetation type in California and current climate change models predict more frequent and severe droughts that could impact plant community structure. Understanding the factors related to species-specific drought mortality is essential to predict such changes. We predicted that life history type, hydraulic traits, and plant size would be related to the ability of species to survive drought. We evaluated the impact of these factors in a mature chaparral stand during the drought of 2014, which has been reported as the most severe in California in the last 1,200 years. We measured tissue water potential, native xylem specific conductivity, leaf specific conductivity, percentage loss in conductivity, and chlorophyll fluorescence for 11 species in February 2014, which was exceptionally dry following protracted drought. Mortality among the 11 dominant species ranged from 0 to 93%. Total stand density was reduced 63.4% and relative dominance of species shifted after the drought. Mortality was negatively correlated with water potential, native xylem specific conductivity, and chlorophyll fluorescence, but not with percent loss in hydraulic conductivity and leaf specific conductivity. The model that best explained mortality included species and plant size as main factors and indicated that larger plants had greater survival for 2 of the species. In general, species with greater resistance to water-stress induced cavitation showed greater mortality levels. Despite adult resprouters typically being more vulnerable to cavitation, results suggest that their more extensive root systems enable them to better access soil moisture and avoid harmful levels of dehydration. These results are consistent with the hypothesis that short-term high intensity droughts have the strongest effect on mature plants of shallow-rooted dehydration tolerant species, whereas deep-rooted dehydration avoiding species fare better in the short-term. Severe droughts can drive changes in chaparral structure as a result of the differential mortality among species.

SummaryThe standard centrifuge method has been frequently used to measure vulnerability to xylem cavitation. This method has recently been questioned. It was hypothesized that open vessels lead to exponential vulnerability curves, which were thought to be indicative of measurement artifact.We tested this hypothesis in stems of olive (Olea europea) because its long vessels were recently claimed to produce a centrifuge artifact. We evaluated three predictions that followed from the open vessel artifact hypothesis: shorter stems, with more open vessels, would be more vulnerable than longer stems; standard centrifuge-based curves would be more vulnerable than dehydration-based curves; and open vessels would cause an exponential shape of centrifuge-based curves.Experimental evidence did not support these predictions. Centrifuge curves did not vary when the proportion of open vessels was altered. Centrifuge and dehydration curves were similar. At highly negative xylem pressure, centrifuge-based curves slightly overestimated vulnerability compared to the dehydration curve. This divergence was eliminated by centrifuging each stem only once.The standard centrifuge method produced accurate curves of samples containing open vessels, supporting the validity of this technique and confirming its utility in understanding plant hydraulics. Seven recommendations for avoiding artefacts and standardizing vulnerability curve methodology are provided.

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.

Plants transport water under negative pressure and this makes their xylem vulnerable to cavitation. Among plant organs, root xylem is often highly vulnerable to cavitation due to water stress. The use of centrifuge methods to study organs, such as roots, that have long vessels are hypothesized to produce erroneous estimates of cavitation resistance due to the presence of open vessels through measured samples. The assumption that roots have long vessels may be premature since data for root vessel length are sparse; moreover, recent studies have not supported the existence of a long-vessel artifact for stems when a standard centrifuge technique was used. We examined resistance to cavitation estimated using a standard centrifuge technique and compared these values with native embolism measurements for roots of seven woody species grown in a common garden. For one species we also measured vulnerability using single-vessel air injection. We found excellent agreement between root native embolism and the levels of embolism measured using a centrifuge technique, and with air-seeding estimates from single-vessel injection. Estimates of cavitation resistance measured from centrifuge curves were biologically meaningful and were correlated with field minimum water potentials, vessel diameter (VD), maximum xylem-specific conductivity (Ksmax) and vessel length. Roots did not have unusually long vessels compared with stems; moreover, root vessel length was not correlated to VD or to the vessel length of stems. These results suggest that root cavitation resistance can be accurately and efficiently measured using a standard centrifuge method and that roots are highly vulnerable to cavitation. The role of root cavitation resistance in determining drought tolerance of woody species deserves further study, particularly in the context of climate change.

The xylem tissue of woody plants performs three principal functions: transport of water, mechanical support of the plant body, and storage of nutrients (Pratt and Jacobsen, 2017). Much previous research has focused on the two functions of water transport and mechanical support in the context of xylem tradeoffs, and there has been less research into the function of storage (however, see Plavcová et al., 2016; Chen et al., 2020). In the present study, we focused on how storage relates to other xylem functions. It is likely that xylem functions are interdependent, such that wood proficient at one function is necessarily poor at another, giving rise to tradeoffs. Tradeoffs can arise due to the structural demands required to support a particular function that is at odds with another function (Janssen et al., 2020). Another possibility is that evolutionary forces lead to wood functions that are associated because they are part of an adaptive suite of traits that determine rates of resource acquisition, use, and turnover (Reich, 2014). One force that strongly influences xylem function is water limitation in seasonal environments. Plants widely differ in dehydration tolerance during drought, which we define as how negative their water potentials are during dry periods. One trait that is associated with dehydration tolerance is cavitation resistance (Pockman and Sperry, 2000; Kursar et al., 2009; Parker et al., 2016). Cavitation occurs when the tension of the xylem sap reaches a point where gas is pulled into water-filled conduits, filling them with gas emboli.

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