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N Michele Holbrook

Publications and source records attributed to N Michele Holbrook.

At least 19 recordsLinked to original sources
PubMed2006

Hydraulic design of pine needles: one-dimensional optimization for single-vein leaves.

Single-vein leaves have the simplest hydraulic design possible, yet even this linear water delivery system can be modulated to improve physiological performance. We determined the optimal distribution of transport capacity that minimizes pressure drop per given investment in xylem permeability along the needle for a given length without a change in total water delivery, or maximizes needle length for a given pressure difference between petiole and needle tip. This theory was tested by comparative analysis of the hydraulic design of three pine species that differ in the length of their needles [Pinus palustris (Engl.) Miller, approximately 50 cm; Pinus ponderosa Lawson & Lawson, approximately 20 cm and Pinus rigida Miller, approximately 5 cm]. In all three species, the distribution of hydraulic permeability was similar to that predicted by the optimum solution. The needles of P. palustris showed an almost perfect match between predicted and actual hydraulic optimum solution, providing evidence that vein design is a significant factor in the hydraulic design of pine leaves.

Pinus↗
PubMed2006

Water relations of baobab trees (Adansonia spp. L.) during the rainy season: does stem water buffer daily water deficits?

Baobab trees are often cited in the literature as water-storing trees, yet few studies have examined this assumption. We assessed the role of stored water in buffering daily water deficits in two species of baobabs (Adansonia rubrostipa Jum. and H. Perrier and Adansonia za Baill.) in a tropical dry forest in Madagascar. We found no lag in the daily onset of sap flow between the base and the crown of the tree. Some night-time sap flow occurred, but this was more consistent with a pattern of seasonal stem water replenishment than with diurnal usage. Intrinsic capacitance of both leaf and stem tissue (0.07-0.08 and 1.1-1.43 MPa(-1), respectively) was high, yet the amount of water that could be withdrawn before turgor loss was small because midday leaf and stem water potentials (WPs) were near the turgor-loss points. Stomatal conductance was high in the daytime but then declined rapidly, suggesting an embolism-avoidance strategy. Although the xylem of distal branches was relatively vulnerable to cavitation (P50: 1.1-1.7 MPa), tight stomatal control and minimum WPs near--1.0 MPa maintained native embolism levels at 30-65%. Stem morphology and anatomy restrict water movement between storage tissues and the conductive pathway, making stored-water usage more appropriate to longer-term water deficits than as a buffer against daily water deficits.

Adansonia↗
PubMed2006

Declining hydraulic efficiency as transpiring leaves desiccate: two types of response.

The conductance of transpiring leaves to liquid water (Kleaf) was measured across a range of steady-state leaf water potentials (Psileaf). Manipulating the transpiration rate in excised leaves enabled us to vary Psileaf in the range -0.1 MPa to less than -1.5 MPa while using a flowmeter to monitor the transpiration stream. Employing this technique to measure how desiccation affects Kleaf in 19 species, including lycophytes, ferns, gymnosperms and angiosperms, we found two characteristic responses. Three of the six angiosperm species sampled maintained a steady maximum Kleaf while Psileaf remained above -1.2 MPa, although desiccation of leaves beyond this point resulted in a rapid decline in Kleaf. In all other species measured, declining Psileaf led to a proportional decrease in Kleaf, such that midday Psileaf of unstressed plants in the field was sufficient to depress Kleaf by an average of 37%. It was found that maximum Kleaf was strongly correlated with maximum CO2 assimilation rate, while Kleaf = 0 occurred at a Psileaf slightly less negative than at leaf turgor loss. A strong linear correlation across species between Psileaf at turgor loss and Psileaf at Kleaf = 0 raises the possibility that declining Kleaf was related to declining cell turgor in the leaf prior to the onset of vein cavitation. The vulnerability of leaves rehydrating after desiccation was compared with vulnerability of leaves during steady-state evaporation, and differences between methods suggest that in many cases vein cavitation occurs only as Kleaf approaches zero.

Carbon Dioxide↗
PubMed2006

Baobab trees (Adansonia) in Madagascar use stored water to flush new leaves but not to support stomatal opening before the rainy season.

Baobab trees (Adansonia, Bombacaceae) are widely thought to store water in their stems for use when water availability is low. We tested this hypothesis by assessing the role of stored water during the dry season in three baobab species in Madagascar. In the dry season, leaves are present only during and after leaf flush. We quantified the relative contributions of stem and soil water during this period through measures of stem water content, sap flow and stomatal conductance. Rates of sap flow at the base of the trunk were near zero, indicating that leaf flushing was almost entirely dependent on stem water. Stem water content declined by up to 12% during this period, yet stomatal conductance and branch sap flow rates remained very low. Stem water reserves were used to support new leaf growth and cuticular transpiration, but not to support stomatal opening before the rainy season. Stomatal opening coincided with the onset of sap flow at the base of the trunk and occurred only after significant rainfall.

Adansonia↗
PubMed2006

Leaf hydraulics.

Leaves are extraordinarily variable in form, longevity, venation architecture, and capacity for photosynthetic gas exchange. Much of this diversity is linked with water transport capacity. The pathways through the leaf constitute a substantial (>or=30%) part of the resistance to water flow through plants, and thus influence rates of transpiration and photosynthesis. Leaf hydraulic conductance (K(leaf)) varies more than 65-fold across species, reflecting differences in the anatomy of the petiole and the venation architecture, as well as pathways beyond the xylem through living tissues to sites of evaporation. K(leaf) is highly dynamic over a range of time scales, showing circadian and developmental trajectories, and responds rapidly, often reversibly, to changes in temperature, irradiance, and water supply. This review addresses how leaf structure and physiology influence K(leaf), and the mechanisms by which K(leaf) contributes to dynamic functional responses at the level of both individual leaves and the whole plant.

Gases↗
PubMed2005

Water stress deforms tracheids peripheral to the leaf vein of a tropical conifer.

Just as a soggy paper straw is prone to yielding under the applied suction of a thirsty drinker, the xylem tracheids in leaves seem prone to collapse as water potential declines, impeding their function. Here we describe the collapse, under tension, of lignified cells peripheral to the leaf vein of a broad-leaved rainforest conifer, Podocarpus grayi de Laub. Leaves of Podocarpus are characterized by an array of cylindrical tracheids aligned perpendicular to the leaf vein, apparently involved in the distribution of water radially through the mesophyll. During leaf desiccation the majority of these tracheids collapsed from circular to flat over the water potential range -1.5 to -2.8 MPa. An increase in the percentage of tracheids collapsed during imposed water stress was mirrored by declining leaf hydraulic conductivity (K(leaf)), implying a direct effect on water transport efficiency. Stomata responded to water stress by closing at -2.0 MPa when 45% of cells were collapsed and K(leaf) had declined by 25%. This was still substantially before the initial indications of cavitation-induced loss of hydraulic conductance in the leaf vein, at -3 MPa. Plants droughted until 49% of tracheids had collapsed were found to fully recover tracheid shape and leaf function 1 week after rewatering. A simple mechanical model of tracheid collapse, derived from the theoretical buckling pressure for pipes, accurately predicted the collapse dynamics observed in P. grayi, substantiating estimates of cell wall elasticity and measured leaf water potential. The possible adaptive advantages of collapsible vascular tissue are discussed.

Biophysical Phenomena↗
PubMed2005

Leaf hydraulic capacity in ferns, conifers and angiosperms: impacts on photosynthetic maxima.

* The hydraulic plumbing of vascular plant leaves varies considerably between major plant groups both in the spatial organization of veins, as well as their anatomical structure. * Five conifers, three ferns and 12 angiosperm trees were selected from tropical and temperate forests to investigate whether the profound differences in foliar morphology of these groups lead to correspondingly profound differences in leaf hydraulic efficiency. * We found that angiosperm leaves spanned a range of leaf hydraulic conductance from 3.9 to 36 mmol m2 s-1 MPa-1, whereas ferns (5.9-11.4 mmol m-2 s-1 MPa-1) and conifers (1.6-9.0 mmol m-2 s-1 MPa-1) were uniformly less conductive to liquid water. Leaf hydraulic conductance (Kleaf) correlated strongly with stomatal conductance indicating an internal leaf-level regulation of liquid and vapour conductances. Photosynthetic capacity also increased with Kleaf, however, it became saturated at values of Kleaf over 20 mmol m-2 s-1 MPa-1. * The data suggest that vessels in the leaves of the angiosperms studied provide them with the flexibility to produce highly conductive leaves with correspondingly high photosynthetic capacities relative to tracheid-bearing species.

Ecosystem↗
PubMed2005

Leaf hydraulic architecture correlates with regeneration irradiance in tropical rainforest trees.

The leaf hydraulic conductance (K(leaf)) is a major determinant of plant water transport capacity. Here, we measured K(leaf), and its basis in the resistances of leaf components, for fully illuminated leaves of five tree species that regenerate in deep shade, and five that regenerate in gaps or clearings, in Panamanian lowland tropical rainforest. We also determined coordination with stomatal characters and leaf mass per area. K(leaf) varied 10-fold across species, and was 3-fold higher in sun- than in shade-establishing species. On average, 12% of leaf hydraulic resistance (= 1/K(leaf)) was located in the petiole, 25% in the major veins, 25% in the minor veins, and 39% outside the xylem. Sun-establishing species had a higher proportion of leaf resistance in the xylem. Across species, component resistances correlated linearly with total leaf resistance. K(leaf) correlated tightly with indices of stomatal pore area, indicating a coordination of liquid- and vapor-phase conductances shifted relative to that of temperate woody species. Leaf hydraulic properties are integrally linked in the complex of traits that define differences in water use and carbon economy across habitats and vegetation zones.

Photobiology↗
PubMed2004

Evolution of xylem lignification and hydrogel transport regulation.

In vascular plants, the polysaccharide-based walls of water-conducting cells are strengthened by impregnation with the polyphenolic polymer lignin. The fine-scale patterning of lignin deposition in water-conducting cells is shown here to vary phylogenetically across vascular plants. The extent to which water transport in xylem cells can be modified in response to changes in the ionic content of xylem sap also is shown to vary in correlation with variation in lignification patterns, consistent with the proposed mechanism for hydraulic response through size change of middle-lamella pectins. This covariation suggests that the fine-scale distribution of hydrophilic polysaccharides and hydrophobic lignin can affect hydraulic as well as mechanical properties, and that the evolutionary diversification of vascular cells thus reflects biochemical as well as morphological innovations evolved to fulfill opposing cell functions of transport and structural support.

Biological Evolution↗
PubMed2004

Functional design space of single-veined leaves: role of tissue hydraulic properties in constraining leaf size and shape.

BACKGROUND AND AIMS: Morphological diversity of leaves is usually quantified with geometrical characters, while in many cases a simple set of biophysical parameters are involved in constraining size and shape. One of the main physiological functions of the leaf is transpiration and thus one can expect that leaf hydraulic parameters can be used to predict potential morphologies, although with the caveat that morphology in turn influences physiological parameters including light interception and boundary layer thickness and thereby heat transfer and net photosynthesis. METHODS: An iterative model was used to determine the relative sizes and shapes that are functionally possible for single-veined leaves as defined by their ability to supply the entire leaf lamina with sufficient water to prevent stomatal closure. The model variables include the hydraulic resistances associated with vein axial and radial transport, as well as with water movement through the mesophyll and the leaf surface. KEY RESULTS: The four parameters included in the model are sufficient to define a hydraulic functional design space that includes all single-veined leaf shapes found in nature, including scale-, awl- and needle-like morphologies. This exercise demonstrates that hydraulic parameters have dissimilar effects: surface resistance primarily affects leaf size, while radial and mesophyll resistances primarily affect leaf shape. CONCLUSIONS: These distinctions between hydraulic parameters, as well as the differential accessibility of different morphologies, might relate to the convergent evolutionary patterns seen in a variety of fossil lineages concerning overall morphology and anatomical detail that frequently have evolved in linear and simple multi-veined leaves.

Models, Biological↗
PubMed2004

Root-gel interactions and the root waving behavior of Arabidopsis.

Arabidopsis roots grown on inclined agarose gels exhibit a sinusoidal growth pattern known as root waving. While root waving has been attributed to both intrinsic factors (e.g. circumnutation) and growth responses to external signals such as gravity, the potential for physical interactions between the root and its substrate to influence the development of this complex phenotype has been generally ignored. Using a rotating stage microscope and time-lapse digital imaging, we show that (1) root tip mobility is impeded by the gel surface, (2) this impedance causes root tip deflections by amplifying curvature in the elongation zone in a way that is distinctly nontropic, and (3) root tip impedance is augmented by normal gravitropic pressure applied by the root tip against the gel surface. Thus, both lateral corrective bending near the root apex and root tip impedance could be due to different vector components of the same graviresponse. Furthermore, we speculate that coupling between root twisting and bending is a mechanical effect resulting from root tip impedance.

Arabidopsis↗
PubMed2004

Changes in pit membrane porosity due to deflection and stretching: the role of vestured pits.

The effect of increasing pressure difference (DeltaP) on intervessel pit membrane porosity was studied in two angiosperm tree species with differing pit architecture. Fraxinus americana L. possesses typical angiosperm bordered pit structure while Sophora japonica L. exhibits well-developed vestures in intervessel pit chambers. It was hypothesized (a) that large DeltaP across intervessel pits would cause the deflection of pit membranes in the stems of F. americana resulting in significant increases in porosity and thus lower cavitation thresholds, and (b) that the presence of vestures would prevent the deflection of pit membranes in S. japonica. To determine if the porosity of pit membranes increased under mechanical stress, suspensions of colloidal gold, 5 nm and 20 nm in diameter, were perfused across intervessel pit membranes at DeltaP ranging from 0.25 MPa to 6.0 MPa. The effect of increasing DeltaP on membrane porosity was also tested by comparing air seeding thresholds (Pa) in stems perfused with water or a solution with lower surface tension. Air seeding and colloidal gold experiments indicated that pit membrane porosity increased significantly with DeltaP in F. americana. In S. japonica, increases in permeability to colloidal gold with DeltaP were small and maximum pore diameters predicted from Pa were independent of DeltaP, suggesting that vestures limited the degree to which the membrane can be deflected from the centre of the pit cavity. This provides the first experimental evidence that vestures reduce the probability of air seeding through pit membranes.

Air↗
PubMed2004

Hydraulic analysis of water flow through leaves of sugar maple and red oak.

Leaves constitute a substantial fraction of the total resistance to water flow through plants. A key question is how hydraulic resistance within the leaf is distributed among petiole, major veins, minor veins, and the pathways downstream of the veins. We partitioned the leaf hydraulic resistance (R(leaf)) for sugar maple (Acer saccharum) and red oak (Quercus rubra) by measuring the resistance to water flow through leaves before and after cutting specific vein orders. Simulations using an electronic circuit analog with resistors arranged in a hierarchical reticulate network justified the partitioning of total R(leaf) into component additive resistances. On average 64% and 74% of the R(leaf) was situated within the leaf xylem for sugar maple and red oak, respectively. Substantial resistance-32% and 49%- was in the minor venation, 18% and 21% in the major venation, and 14% and 4% in the petiole. The large number of parallel paths (i.e. a large transfer surface) for water leaving the minor veins through the bundle sheath and out of the leaf resulted in the pathways outside the venation comprising only 36% and 26% of R(leaf). Changing leaf temperature during measurement of R(leaf) for intact leaves resulted in a temperature response beyond that expected from changes in viscosity. The extra response was not found for leaves with veins cut, indicating that water crosses cell membranes after it leaves the xylem. The large proportion of resistance in the venation can explain why stomata respond to leaf xylem damage and cavitation. The hydraulic importance of the leaf vein system suggests that the diversity of vein system architectures observed in angiosperms may reflect variation in whole-leaf hydraulic capacity.

Acer↗
PubMed2004

A potential role for xylem-phloem interactions in the hydraulic architecture of trees: effects of phloem girdling on xylem hydraulic conductance.

We investigated phloem-xylem interactions in Acer rubrum L. and Acer saccharum Marsh. Our experimental method allowed us to determine xylem conductance of an intact branch by measuring the flow rate of water supplied at two delivery pressures to the cut end of a small side branch. We found that removal of bark tissue (phloem girdling) upstream of the point at which deionized water was delivered to the branch resulted in a decrease (24% for A. rubrum and 15% for A. saccharum) in branch xylem hydraulic conductance. Declines in hydraulic conductance with girdling were accompanied by a decrease in the osmotic concentration of xylem sap. The decrease in xylem sap concentration following phloem girdling suggests that ion redistribution from the phloem was responsible for the observed decline in hydraulic conductance. When the same measurements were made on branches perfused with KCl solution (approximately 140 mOsm kg(-1)), phloem girdling had no effect on xylem hydraulic conductance. These results suggest a functional link between phloem and xylem hydraulic systems that is mediated by changes in the ionic content of the cell sap.

Acer↗
PubMed2003

Ionic control of the lateral exchange of water between vascular bundles in tomato.

Ions can enhance water flow through the xylem via changes in the hydraulic resistance at border pit membranes. Because flow between adjacent xylem vessels occurs primarily via bordered pit fields, it is hypothesized that xylem sap ion concentrations would affect lateral movement of water more than longitudinal flow. Using tomato as a model system, evidence is presented for ion-mediated changes in xylem hydraulic resistance and the lateral transport of water. Water flow between adjacent xylem bundles increased by approximately 50% in the presence of ions while longitudinal flow only increased by approximately 20%. However, the enhancement of lateral exchange due to ions was magnified by the presence of a pressure difference between vascular bundles. These results indicate that the degree of nutrient-sharing among sectors of a plant may depend on both nutrient concentration and the availability of water in the root zone.

Biological Transport↗
PubMed2003

Vulnerability of xylem vessels to cavitation in sugar maple. Scaling from individual vessels to whole branches.

The relation between xylem vessel age and vulnerability to cavitation of sugar maple (Acer saccharum Marsh.) was quantified by measuring the pressure required to force air across bordered pit membranes separating individual xylem vessels. We found that the bordered pit membranes of vessels located in current year xylem could withstand greater applied gas pressures (3.8 MPa) compared with bordered pit membranes in vessels located in older annular rings (2.0 MPa). A longitudinal transect along 6-year-old branches indicated that the pressure required to push gas across bordered pit membranes of current year xylem did not vary with distance from the growing tip. To understand the contribution of age-related changes in vulnerability to the overall resistance to cavitation, we combined data on the pressure thresholds of individual xylem vessels with measurements of the relative flow rate through each annual ring. The annual ring of the current year contributed only 16% of the total flow measured on 10-cm-long segments cut from 6-year-old branches, but it contributed more than 70% of the total flow when measured through 6-year-old branches to the point of leaf attachment. The vulnerability curve calculated using relative flow rates measured on branch segments were similar to vulnerability curves measured on 6-year-old branches (pressure that reduces hydraulic conductance by 50% = 1.6-2.4 MPa), whereas the vulnerability curve calculated using relative flow rates measured on 6-year-old branches were similar to ones measured on the extension growth of the current year (pressure that reduces hydraulic conductance by 50% = 3.8 MPa). These data suggest that, in sugar maple, the xylem of the current year can withstand larger xylem tensions than older wood and dominates water delivery to leaves.

Acer↗
PubMed2003

Stomatal closure during leaf dehydration, correlation with other leaf physiological traits.

The question as to what triggers stomatal closure during leaf desiccation remains controversial. This paper examines characteristics of the vascular and photosynthetic functions of the leaf to determine which responds most similarly to stomata during desiccation. Leaf hydraulic conductance (K(leaf)) was measured from the relaxation kinetics of leaf water potential (Psi(l)), and a novel application of this technique allowed the response of K(leaf) to Psi(l) to be determined. These "vulnerability curves" show that K(leaf) is highly sensitive to Psi(l) and that the response of stomatal conductance to Psi(l) is closely correlated with the response of K(leaf) to Psi(l). The turgor loss point of leaves was also correlated with K(leaf) and stomatal closure, whereas the decline in PSII quantum yield during leaf drying occurred at a lower Psi(l) than stomatal closure. These results indicate that stomatal closure is primarily coordinated with K(leaf). However, the close proximity of Psi(l) at initial stomatal closure and initial loss of K(leaf) suggest that partial loss of K(leaf) might occur regularly, presumably necessitating repair of embolisms.

Desiccation↗