Brett A. Huggett

Cutting xylem under tension or supersaturated with gas can generate PLC and the appearance of rapid recovery from embolism

We investigated the common assumption that severing stems and petioles under water preserves the hydraulic continuity in the xylem conduits opened by the cut when the xylem is under tension. In red maple and white ash, higher percent loss of conductivity (PLC) in the afternoon occurred when the measurement segment was excised under water at native xylem tensions, but not when xylem tensions were relaxed prior to sample excision. Bench drying vulnerability curves in which measurement samples were excised at native versus relaxed tensions showed a dramatic effect of cutting under tension in red maple, a moderate effect in sugar maple, and no effect in paper birch. We also found that air injection of cut branches (red and sugar maple) at pressures of 0.1 and 1.0 MPa resulted in PLC greater than predicted from vulnerability curves for samples cut 2 min after depressurization, with PLC returning to expected levels for samples cut after 75 min. These results suggest that sampling methods can generate PLC patterns indicative of repair under tension by inducing a degree of embolism that is itself a function of xylem tensions or supersaturation of dissolved gases (air injection) at the moment of sample excision. Implications for assessing vulnerability to cavitation and levels of embolism under field conditions are discussed.

Distribution and mixing of old and new nonstructural carbon in two temperate trees

We know surprisingly little about whole-tree nonstructural carbon (NSC; primarily sugars and starch) budgets. Even less well understood is the mixing between recent photosynthetic assimilates (new NSC) and previously stored reserves. And, NSC turnover times are poorly constrained. We characterized the distribution of NSC in the stemwood, branches, and roots of two temperate trees, and we used the continuous label offered by the radiocarbon (carbon-14, 14C) bomb spike to estimate the mean age of NSC in different tissues. NSC in branches and the outermost stemwood growth rings had the 14C signature of the current growing season. However, NSC in older aboveground and belowground tissues was enriched in 14C, indicating that it was produced from older assimilates. Radial patterns of 14C in stemwood NSC showed strong mixing of NSC across the youngest growth rings, with limited ‘mixing in’ of younger NSC to older rings. Sugars in the outermost five growth rings, accounting for two-thirds of the stemwood pool, had a mean age < 1 yr, whereas sugars in older growth rings had a mean age > 5 yr. Our results are thus consistent with a previously-hypothesized two-pool (‘fast’ and ‘slow’ cycling NSC) model structure. These pools appear to be physically distinct.

Cell longevity and sustained primary growth in palm stems

Longevity, or organismal life span, is determined largely by the period over which constituent cells can function metabolically. Plants, with modular organization (the ability continually to develop new organs and tissues) differ from animals, with unitary organization (a fixed body plan), and this difference is reflected in their respective life spans, potentially much longer in plants than animals. We draw attention to the observation that palm trees, as a group of monocotyledons without secondary growth comparable to that of lignophytes (plants with secondary growth from a bifacial cambium), retain by means of sustained primary growth living cells in their trunks throughout their organismal life span. Does this make palms the longest-lived trees because they can grow as individuals for several centuries? No conventional lignophyte retains living metabolically active differentiated cell types in its trunk for this length of time, even though the tree as a whole can exist for millennia. Does this contrast also imply that the long-lived cells in a palm trunk have exceptional properties, which allows this seeming immortality? We document the long-life of many tall palm species and their inherent long-lived stem cell properties, comparing such plants to conventional trees. We provide a summary of aspects of cell age and life span in animals and plants. Cell replacement is a feature of animal function, whereas conventional trees rely on active growth centers (meristems) to sustain organismal development. However, the long persistence of living cells in palm trunks is seen not as evidence for unique metabolic processes that sustain longevity, but is a consequence of unique constructional features. This conclusion suggests that the life span of plant cells is not necessarily genetically determined.

Aspects of Vessel Dimensions in the Aerial Roots of Epiphytic Araceae

We measured vessel dimensions, most significantly vessel length, in the aerial roots of four epiphytic aroids using a digital camera to photograph sequential sections. Pendulous aerial roots in Araceae can grow from the forest canopy and so reach considerable length (>30 m) before they contact the ground, branch, and become anchored. In the free‐hanging state, the length over which tissue maturation occurs can exceed 1 m. We show that the distinctive medullary vessels do not anastomose and each series of vessels, end to end here termed a “pipe,” must differentiate without interruption throughout the length of the root and do not become fully functional until the ground is reached. Measurements show different vessel parameters, including vessel overlap at each vessel end, which is not usually considered in estimates of hydraulic conductivity. Our method of measurement is simple and direct and shows the topographic relation of all vessels in a single organ, suggesting that vessels in long plant organs can be measured precisely, giving results of value in considering the hydraulic properties of xylem elements.

The role of leaf hydraulic conductance dynamics on the timing of leaf senescence

We tested the hypothesis that an age-dependent reduction in leaf hydraulic conductance (Kleaf) influences the timing of leaf senescence via limitation of the stomatal aperture on xylem compound delivery to leaves of tomato (Solanum lycopersicum L.), the tropical trees Anacardium excelsum Kunth, Pittoniotis trichantha Griseb, and the temperate trees Acer saccharum Marsh. and Quercus rubra L. The onset of leaf senescence was preceded by a decline in Kleaf in tomato and the tropical trees, but not in the temperate trees. Age-dependent changes in Kleaf in tomato were driven by a reduction in leaf vein density without a proportional increase in the xylem hydraulic supply. A decline in stomatal conductance accompanied Kleaf reduction with age in tomato but not in tropical and temperate tree species. Experimental manipulations that reduce the flow of xylem-transported compounds into leaves with open stomata induced early leaf senescence in tomato and A. excelsum, but not in P. trichantha, A. saccharum and Q. rubra leaves. We propose that in tomato, a reduction in Kleaf limits the delivery of xylem-transported compounds into the leaves, thus making them vulnerable to senescence. In the tropical evergreen tree A. excelsum, xylem-transported compounds may play a role in signalling the timing of senescence but are not under leaf hydraulic regulation; leaf senescence in the deciduous trees A. trichanta, A. saccharum and Q. rubra is not influenced by leaf vascular transport.

MicroCT imaging as a tool to study vessel endings in situ

PREMISE OF THE STUDY Despite the strong influence of the frequency and distribution of vessel endings on both hydraulic safety and efficiency, detailed anatomical descriptions or measurements of these structures are generally lacking. METHODS Here we used high-resolution x-ray microcomputed tomography (microCT) to identify and describe xylem vessel endings within Acer rubrum root segments (1.0-2.1 mm diameter, ∼2 mm long). We then compared vessel-lumen diameter, pit density, vessel element length, and perforation plate angle between non-ending vessels (those that traverse an entire segment) and those that end within a segment using three-dimensional image analysis. KEY RESULTS We found 214 vessel endings, 37 complete vessels, and 385 non-ending vessels within four A. rubrum root segments. Vessels that ended within the segments tended to have more acute perforation plate angles and had a smaller diameter than those that did not end within the segments. Most vessel diameters tapered within the last few vessel elements, but the perforation plate angle apparently changed over longer distances. Intervessel pit density and vessel element length did not differ between ending and non-ending vessels. CONCLUSIONS Vessel endings were surprisingly frequent in A. rubrum roots despite the common perception that root vessels are longer than vessels in other tissues. MicroCT proved to be a useful tool for studying the three-dimensional arrangement of vessel endings within xylem networks, and these data will be helpful in developing a better understanding of vessel ending microstructure and function.

Stem anatomy in the spiny american palm Bactris (Arecaceae-Bactridinae)

Bactris trunks are made entirely by long sustained primary growth so that mechanical properties develop progressively as the trunk matures. Anatomical features vary throughout the stem so that one small sample is a limited representation. This limits the use for stem histology with systematic purposes. We examined stem anatomy in 17 out of 73 species representing all major sub-generic groups. Stem features do not associate well with the subdivision of the genus. Distinctive structural and developmental features: are the presence of fiber strands independent of the vascular tissues, the modified ground parenchyma with a late formation of air-lacunae, and the early inception of the mechanically significant outer central cylinder. Cell expansion and fiber wall thickening develop as the stem ages, but vary in their expression in stems of different diameter. Our example thus provides an explanation for the difficulty of identifying the systematic position of fossil palm stems.

Whole-tree nonstructural carbohydrate storage and seasonal dynamics in five temperate species

Summary Despite the importance of nonstructural carbohydrates (NSC) for growth and survival in woody plants, we know little about whole‐tree NSC storage. The conventional theory suggests that NSC reserves will increase over the growing season and decrease over the dormant season. Here, we compare storage in five temperate tree species to determine the size and seasonal fluctuation of whole‐tree total NSC pools as well as the contribution of individual organs. NSC concentrations in the branches, stemwood, and roots of 24 trees were measured across 12 months. We then scaled up concentrations to the whole‐tree and ecosystem levels using allometric equations and forest stand inventory data. While whole‐tree total NSC pools followed the conventional theory, sugar pools peaked in the dormant season and starch pools in the growing season. Seasonal depletion of total NSCs was minimal at the whole‐tree level, but substantial at the organ level, particularly in branches. Surprisingly, roots were not the major storage organ as branches stored comparable amounts of starch throughout the year, and root reserves were not used to support springtime growth. Scaling up NSC concentrations to the ecosystem level, we find that commonly used, process‐based ecosystem and land surface models all overpredict NSC storage.

A Novel Type of Fiber in the Leaves of the Cycad Dioon

Premise of research. The extant genera of cycads (order Cycadales) can be readily distinguished by the anatomy of their leaflets. In particular, the genus Dioon possesses a unique cellulosic fiber type in the leaflet mesophyll, not found in any other genus. We examine living material of all 15 species of Dioon to confirm preliminary observations and provide details of wall structure in comparison with fibers in the leaf axis. Methodology. Unembedded sample material was sectioned on a sliding microtome transversely and longitudinally, and standard histochemical tests were utilized to identify cell types. SEM was utilized to analyze fiber anatomy. Maceration of samples provided cell types for comparison. Pivotal results. All species of Dioon have the same configuration of cell types in the leaflets, the most conspicuous element being elongated fibers with multilamellate cellulosic cell walls. Such fibers do not appear in the leaf axis. This fiber type is seemingly unique among all cycad genera. Conclusions. Dioon leaflets possess a fiber type of distinctive wall structure, unique among the extant cycads. This finding clarifies an earlier report in which such fibers were described as “gelatinous fibers.”

Impact of hemlock woolly adelgid (Adelges tsugae) infestation on xylem structure and function and leaf physiology in eastern hemlock (Tsuga canadensis)

Hemlock woolly adelgid (Adelges tsugae Annand) (HWA) is an invasive insect that feeds upon the foliage of eastern hemlock (Tsuga canadensis (L.) Carrière) trees, leading to a decline in health and often mortality. The exact mechanism leading to the demise of eastern hemlocks remains uncertain because little is known about how HWA infestation directly alters the hosts physiology. To evaluate the physiological responses of eastern hemlock during early infestation of HWA, we measured needle loss, xylem hydraulic conductivity, vulnerability to cavitation, tracheid anatomy, leaf-level gas exchange, leaf water potential and foliar cation and nutrient levels on HWA-infested and noninfested even-aged trees in an experimental garden. HWA infestation resulted in higher xylem hydraulic conductivity correlated with an increase in average tracheid lumen area and no difference in vulnerability to cavitation, indicating that needle loss associated with HWA infestation could not be attributed to reduced xylem transport capacity. HWA-infested trees exhibited higher rates of net photosynthesis and significant changes in foliar nutrient partitioning, but showed no differences in branch increment growth rates compared with noninfested trees. This study suggests that HWA-induced decline in the health of eastern hemlock trees is not initially caused by compromised water relations or needle loss.