Formation and Stabilization of Rhizosheaths of Zea mays L. (Effect of Soil Water Content)

Field observations have shown that rhizosheaths of grasses formed under dry conditions are larger, more coherent, and more strongly bound to the roots than those formed in wet soils. We have quantified these effects in a model system in which corn (Zea mays L.) primary roots were grown through a 30-cm-deep prepared soil profile that consisted of a central, horizontal, "dry" (9% water content) or "wet" (20% water content) layer (4 cm thick) sandwiched between damp soil (15-17% water content). Rhizosheaths formed in dry layers were 5 times the volume of the subtending root. In wet layers, rhizosheaths were only 1.5 times the root volume. Fractions of the rhizosheath soil were removed from individual roots by three successive treatments; sonication, hot water, and abrasion. Sonication removed 50 and 90% of the soil from rhizosheaths formed in dry and wet soils, respectively. After the heat treatment, 35% of the soil still adhered to those root portions where rhizosheaths had developed in dry soil, compared with 2% where sheaths had formed in wet soil. Root hairs were 4.5 times more abundant and were more distorted on portions of roots from dry layers than from wet layers. Drier soil enhanced adhesiveness of rhizosheath mucilages and stimulated the formation of root hairs; both effects stabilize the rhizosheath. Extensive and stable rhizosheaths may function in nutrient acquisition in dry soils.     

Aluminium tolerance of root hairs underlies genotypic differences in rhizosheath size of wheat (Triticum aestivum) grown on acid soil

We found significant genetic variation in the ability of wheat (Triticum aestivum) to form rhizosheaths on acid soil and assessed whether differences in aluminium (Al(3+) ) tolerance of root hairs between genotypes was the physiological basis for this genetic variation. A method was developed to rapidly screen rhizosheath size in a range of wheat genotypes. Backcrossed populations were generated from cv Fronteira (large rhizosheath) using cv EGA-Burke (small rhizosheath) as the recurrent parent. A positive correlation existed between rhizosheath size on acid soil and root hair length. In hydroponic experiments, root hairs of the backcrossed lines with large rhizosheaths were more tolerant of Al(3+) toxicity than the backcrossed lines with small rhizosheaths. We conclude that greater Al(3+) tolerance of root hairs underlies the larger rhizosheath of wheat grown on acid soil. Tolerance of the root hairs to Al(3+) was largely independent of the TaALMT1 gene which suggests that different genes encode the Al(3+) tolerance of root hairs. The maintenance of longer root hairs in acid soils is important for the efficient uptake of water and nutrients.     

Small differences in root distributions allow resource niche partitioning

1. Deep roots have long been thought to allow trees to coexist with shallow‐rooted grasses. However, data demonstrating how root distributions affect water uptake and niche partitioning are uncommon.
2. We describe tree and grass root distributions using a depth‐specific tracer experiment six times over two years in a subtropical savanna, Kruger National Park, South Africa. These point‐in‐time measurements were then used in a soil water flow model to simulate continuous water uptake by depth and plant growth form (trees and grasses) across two growing seasons. This allowed estimates of the total amount of water a root distribution could absorb as well as the amount of water a root distribution could absorb in excess of the other rooting distribution (i.e., unique hydrological niche).
3. Most active tree and grass roots were in shallow soils: The mean depth of water uptake was 22 cm for trees and 17 cm for grasses. Slightly deeper rooting distributions provided trees with 5% more soil water than the grasses in a drier season, but 13% less water in a wetter season. Small differences also provided each rooting distribution (tree or grass) with unique hydrological niches of 4 to 13 mm water.
4. The effect of rooting distributions has long been inferred. By quantifying the depth and timing of water uptake, we demonstrated how even small differences in rooting distributions can provide plants with resource niches that can contribute to species coexistence. Differences in total water uptake and unique hydrological niche sizes were small in this system, but they indicated that tradeoffs in rooting strategies can be expected to contribute to tree and grass coexistence because 1) competitive advantages change over time and 2) plant growth forms always have access to a soil resource pool that is not available to the other plant growth form.

     

Ecosystem wicks: Woodland trees enhance water infiltration in a fragmented agricultural landscape in eastern Australia

Abstract Since European settlement, Eucalyptus box woodlands have been substantially modified by agricultural practices, and in many areas in southern Australia are now restricted to scattered or clumped trees. We report here on a study to examine the impact of trees on water flow (infiltration) in an agricultural landscape with substantial areas of extant native vegetation. We examined infiltration through coarse‐ and fine‐textured soils within four landscape strata, the zones below Eucalyptus melliodora and Callitris glaucophylla canopies, the intertree zone dominated by perennial grasses and a landscape homogenized by cultivation and dominated by annual crops. We measured sorptivity, the early phase of water flow, and steady‐state infiltration with disc permeameters at two supply potentials. These different potentials enabled us to separate infiltration into (i) flow through large (biopores) and small pores and (ii) flow through small pores only where biopores are prevented from conducting water. On the fine‐textured soils, both sorptivity and steady‐state infiltration were significantly greater (approximately fivefold) under the timbered strata compared with the grassy slopes or cultivation. Differences were attributable to the greater proportion of macropores below the tree canopies compared with the nontimbered strata. The lack of a significant difference on the coarse‐textured soils, despite their macropore status, was attributed to differences in surface litter and plant cover, which would maintain continuous macropores at the surface and thus conduct large amounts of water. The tendency of slopes covered by cryptogamic crusts and grasses to shed run‐off and for the trees to absorb substantial quantities of water reinforced the important ecological service provided by trees, which moderates large run‐off events and captures small amounts of water leaking from the grassy patches. In the absence of these ‘ecosystem wicks’, run‐off would find its way into regional groundwater and contribute to rising salinity.     

Root niche partitioning among grasses, saplings, and trees measured using a tracer technique

Niche partitioning of resources by plants is believed to be a fundamental aspect of plant coexistence and biogeochemical cycles; however, measurements of the timing and location of resource use are often lacking because of the difficulties of belowground research. To measure niche partitioning of soil water by grasses, planted saplings, and trees in a mesic savanna (Kruger National Park, South Africa), we injected deuterium oxide into 102,000 points in 15, 154-m² plots randomly assigned to one of five depths (0–120 cm) and one of three time periods during the 2008/2009 growing season. Grasses, saplings and trees all demonstrated an exponential decline in water uptake early in the season when resources were abundant. Later in the season, when resources were scarce, grasses continued to extract the most water from the shallowest soil depths (5 cm), but saplings and trees shifted water uptake to deeper depths (30–60 cm). Saplings, in particular, rapidly established roots to at least 1 m and used these deep roots to a greater extent than grasses or trees. Helping to resolve contradictory observations of the relative importance of deep and shallow roots, our results showed that grasses, saplings and trees all extract the most water from shallow soils when it is available but that woody plants can rapidly shift water uptake to deeper soils when resources are scarce. Results highlight the importance of temporal changes in water uptake and the problems with inferring spatial and temporal partitioning of soil water uptake from root biomass measurements alone.     

Spatial variations of microinvertebrates across different microhabitats of temporary floodplains of lower Okavango Delta,Botswana

Temporary floodplains of the Okavango Delta have been suggested as important nursery sites for juvenile fish. Fish production in the Delta is dependent on how well temporary floodplains are maintained. A study was conducted to understand how floodplains function so as to put measures in place that will sustain temporary floodplains in a high productive state. During high floods, distinct microhabitats (open water, sedges and inundated terrestrial grasses) develop in which microinvertebrate community structure was assessed across such habitats during the 2009–2010 flooding season. Depth was highest (ANOVA, P < 0.05) in the open‐water microhabitat and lowest within inundated grasses. Conductivity, total phosphorus and chlorophyll a were all highest (ANOVA, P < 0.05) within inundated grasses than in other microhabitats. A total of 46 microinvertebrate taxa were identified, and they were most diverse within the sedges. The abundance of microinvertebrates was highest (ANOVA, P < 0.05) within the inundated grasses and lowest within the open‐water zone. Given that microinvertebrates are an important food source for juvenile fish, their presence in high densities within inundated terrestrial grasses is a key determinant of fish production. This suggests the need for serious consideration of the importance of temporary floodplains when formulating conservation strategies for the Okavango Delta.     

Nitrogen Fixation Associated with Sand Grain Root Sheaths (Rhizosheaths) of Certain Xeric Grasses

Nitrogen fixation (acetylene reduction) was found to be associated with sand grain root sheaths (rhizoseaths) occurring on the following xeric grasses: Oryzopsis hymenoides (Roem. and Shult.) Ricker, Agropyron dasystachyum (Hook.) Scrib., Stipa comata Trin. and Rupr., and Aristida purpurea Nutt. Acetylene reduction rates associated with whole plant specimens of these species varied from 515 to 920 nmol C₂H₄/(g dry wt.) × (6 days). Nitrogenase activity was shown to be associated with the rhizosheaths. Bacillus polymyxa-like nitrogen fixers were isolated from the rhizosheaths of each grass. The isolates reduced acetylene and assimilated ¹⁵N₂.     

Understanding the Hydraulics of Porous Pipes: Tradeoffs Between Water Uptake and Root Length Utilization

The water uptake region in roots is several hundred times longer than the root diameter. The distributed nature of the uptake zone requires that the hydraulic design of roots be understood by analogy to flow through a “porous pipe.” Here we present results of an analytical and experimental investigation that allowed an in-depth analysis of root hydraulic properties. Measurements on nodal maize roots confirm the nonlinear distribution of water uptake predicted by the porous pipe model. The major design parameter governing the distribution of water uptake along a porous pipe is the ratio between its axial and radial hydraulic resistance. However, total flow is proportional to the pipe's overall resistance. These results suggest the existence of a tradeoff between the effective utilization of root length and the total capacity for water uptake.     

Exploring the structural changes in nitrogen-fixing microorganisms of rhizosheath during the growth of Stipagrostis pennata in the desert

Purpose: Rhizosheath is an adaptive feature for the survival of Stipagrostis pennata in desert systems. Although microorganisms play important ecological roles in promoting the nitrogen cycle of rhizosheath, the diversity and function of nitrogen-fixing microorganism communities have not been fully understood.Materials and methods: Therefore, the aim of the present study is to explore the nitrogen fixation ability of rhizosheaths and the changes in abundance of nitrogen-fixing microorganisms at different growth periods of S. pennata. We sequenced the nifH gene through sequencing to identify the structure and diversity of nitrogen-fixing microorganisms of S. pennata at different growth periods of rhizosheaths.Results: A total of 1256 operational taxonomic units (OTUs) were identified by nifH sequence and distributed in different growth periods. There were five OTUs distributed in each sample at the same time, and the abundance and diversity of microorganisms in fruit period were much higher than those in other periods. Mainly four phyla were involved, among which Proteobacteria was the most abundant in all groups.Conclusions: In general, the present study investigated the abundance and characteristics of nitrogen-fixing microorganisms of rhizosheaths in different growth periods of S. pennata. It also may elucidate and indicate that the structure of nitrogen-fixing microorganisms of rhizosheaths in different growth periods of S. pennata had changed.     

Drivers of Variation in Aboveground Net Primary Productivity and Plant Community Composition Differ Across a Broad Precipitation Gradient

Aboveground net primary production (ANPP) is a key integrator of C uptake and energy flow in many terrestrial ecosystems. As such, ecologists have long sought to understand the factors driving variation in this important ecosystem process. Although total annual precipitation has been shown to be a strong predictor of ANPP in grasslands across broad spatial scales, it is often a poor predictor at local scales. Here we examine the amount of variation in ANPP that can be explained by total annual precipitation versus precipitation during specific periods of the year (precipitation periods) and nutrient availability at three sites representing the major grassland types (shortgrass steppe, mixed-grass prairie, and tallgrass prairie) spanning the broad precipitation gradient of the U.S. Central Great Plains. Using observational data, we found that precipitation periods and nutrient availability were much stronger predictors of site-level ANPP than total annual precipitation. However, the specific nutrients and precipitation periods that best predicted ANPP differed among the three sites. These effects were mirrored experimentally at the shortgrass and tallgrass sites, with precipitation and nutrient availability co-limiting ANPP, but not at the mixed-grass site, where nutrient availability determined ANPP exclusive of precipitation effects. Dominant grasses drove the ANPP response to increased nutrient availability at all three sites. However, the relative responses of rare grasses and forbs were greater than those of the dominant grasses to experimental nutrient additions, thus potentially driving species turnover with chronic nutrient additions. This improved understanding of the factors driving variation in ANPP within ecosystems spanning the broad precipitation gradient of the Great Plains will aid predictions of alterations in ANPP under future global change scenarios.     

Spatial partitioning of the soil water resource between grass and shrub components in a West African humid savanna

Most savanna water balance models assume water partitioning between grasses and shrubs in a two-layer hypothesis, but this hypothesis has not been tested for humid savanna environments. Spatial partitioning of soil water between grasses and shrubs was investigated in a West African humid savanna by comparing the isotopic composition (oxygen-18 and deuterium) of soil water and plant stem water during rainy and dry conditions. Both grass and shrub species acquire most of their water from the top soil layer during both rainy and dry periods. A shift of water uptake pattern towards deeper horizons was observed only at the end of the dry season after shrub defoliation. The mean depth of water uptake, as determined by the isotopic signature of stem water, was consistent with grass and shrub root profiles and with changes in soil water content profiles as surveyed by a neutron probe. This provides evidence for potentially strong competition between shrubs and grasses for soil water in these humid savannas. Limited nutrient availability may explain these competitive interactions. These results enhance our understanding of shrub-grass interactions, and will contribute to models of ecosystem functioning in humid savannas.     

Revisiting the Two-Layer Hypothesis: Coexistence of Alternative Functional Rooting Strategies in Savannas

The two-layer hypothesis of tree-grass coexistence posits that trees and grasses differ in rooting depth, with grasses exploiting soil moisture in shallow layers while trees have exclusive access to deep water. The lack of clear differences in maximum rooting depth between these two functional groups, however, has caused this model to fall out of favor. The alternative model, the demographic bottleneck hypothesis, suggests that trees and grasses occupy overlapping rooting niches, and that stochastic events such as fires and droughts result in episodic tree mortality at various life stages, thus preventing trees from otherwise displacing grasses, at least in mesic savannas. Two potential problems with this view are: 1) we lack data on functional rooting profiles in trees and grasses, and these profiles are not necessarily reflected by differences in maximum or physical rooting depth, and 2) subtle, difficult-to-detect differences in rooting profiles between the two functional groups may be sufficient to result in coexistence in many situations. To tackle this question, I coupled a plant uptake model with a soil moisture dynamics model to explore the environmental conditions under which functional rooting profiles with equal rooting depth but different depth distributions (i.e., shapes) can coexist when competing for water. I show that, as long as rainfall inputs are stochastic, coexistence based on rooting differences is viable under a wide range of conditions, even when these differences are subtle. The results also indicate that coexistence mechanisms based on rooting niche differentiation are more viable under some climatic and edaphic conditions than others. This suggests that the two-layer model is both viable and stochastic in nature, and that a full understanding of tree-grass coexistence and dynamics may require incorporating fine-scale rooting differences between these functional groups and realistic stochastic climate drivers into future models.     

Temperature and functional traits influence differences in nitrogen uptake capacity between native and invasive grasses

Performance differences between native and exotic invasive plants are often considered static, but invasive grasses may achieve growth advantages in western North America shrublands and steppe under only optimal growing conditions. We examine differences in N uptake and several morphological variables that influence uptake at temperatures between 5 and 25 °C. We contrast two native perennial grasses in western North America: Elymus elymoides and Pseudoroegneria spicata; two invasive annual grasses: Bromus tectorum and Taeniatherum caput-medusae; and one highly selected non-native perennial grass: Agropyron cristatum. The influence of temperature on N uptake is poorly characterized, yet these invasive annual grasses are known to germinate in warm soils in the autumn, and both experience cool soils during the short growing season following snowmelt in the spring. To further explore the influence of temperature on the correlation between morphological variables and N uptake, our data are applied to a previously published path model and one proposed here. Differences in N uptake between native and invasive grasses were small at the lowest temperature, but were large at the highest temperature. At lower temperatures, uptake of N by annuals and perennials was correlated with leaf N and mass. At higher temperatures, uptake by annuals was correlated only with these leaf traits, but uptake by perennials was correlated with these leaf traits as well as root N and mass. Consequently, our results imply that annual grasses face fewer morphological constraints on N uptake than perennial grasses, and annual grasses may gain further advantage in warmer temperature conditions or during more frequent warm periods.     

Resistance to Water Flow in the Seminal Roots of Wheat

Water uptake by the seminal root system of wheat is describedby a model which incorporates radial and axial resistance toflow in the root and the resistance to flow in the soil. Inan experiment where wheat plants were dependent on subsoil waterextracted from below 45 cm by one, three, or five seminal roots,calculations indicated that the axial resistance was the dominantresistance within the soil-root system and that flow rate changedpredictably with changing axial resistance. The model also indicatedthat the observed shape of the metaxylem vessel in the seminalaxes has important effects on the pattern of water uptake.     

Variation in root system traits among African semi‐arid savanna grasses: Implications for drought tolerance

In arid to semi‐arid grasslands and savannas, plant growth, population dynamics, and productivity are consistently and strongly limited by soil water and nutrient availability. Adaptive traits of the root systems of grasses in these ecosystems are crucial to their ability to cope with strong water and/or nutrient limitation and the increasing drought stress associated with ecosystem degradation or projected climate change. We studied 18 grass species in semi‐arid savanna of the Kalahari region of Botswana to quantify interspecific variation in three important root system traits including root system architecture, rhizosheath thickness and mycorrhizal colonization. Drought‐tolerant species and shorter‐lived species showed greater rhizosheath thickness and fine root development but lower mycorrhizal colonization compared to later successional climax grasses and those characteristic of wetter sites. In addition, there was a significant positive correlation between root fibrousness index and rhizosheath thickness among species and a weak negative correlation between root fibrousness index and mycorrhizal colonization. These patterns suggest that an extensive fine root system and rhizosheath development may be important complementary traits of grasses coping with drought conditions, the former aiding in the acquisition of water by the grass plant and the latter aiding in water uptake and retention, and reducing water loss in the rhizosphere. Within species, both rhizosheath development and mycorrhizal colonization were significantly greater in a wet year than in a year with below‐average precipitation. The observed patterns suggest that the primary benefit of rhizosheath development in African savanna grasses is improved drought tolerance and that it is a plastic trait that can be adjusted annually to changing environmental conditions. The functioning of mycorrhizal symbiosis is likely to be relatively more important in infertile savannas where nutrient limitation is higher relative to water limitation.     

Zooming in on coarse plant functional types—simulated response of savanna vegetation composition in response to aridity and grazing

Precipitation and land use in terms of livestock grazing have been identified as two of the most important drivers structuring the vegetation composition of semi-arid and arid savannas. Savanna research on the impact of these drivers has widely applied the so-called plant functional type (PFT) approach, grouping the vegetation into two or three broad types (here called meta-PFTs): woody plants and grasses, which are sometimes divided into perennial and annual grasses. However, little is known about the response of functional traits within these coarse types towards water availability or livestock grazing. In this study, we extended an existing eco-hydrological savanna vegetation model to capture trait diversity within the three broad meta-PFTs to assess the effects of both grazing and mean annual precipitation (MAP) on trait composition along a gradient of both drivers. Our results show a complex pattern of trait responses to grazing and aridity. The response differs for the three meta-PFTs. From our findings, we derive that trait responses to grazing and aridity for perennial grasses are similar, as suggested by the convergence model for grazing and aridity. However, we also see that this only holds for simulations below a MAP of 500 mm. This combined with the finding that trait response differs between the three meta-PFTs leads to the conclusion that there is no single, universal trait or set of traits determining the response to grazing and aridity. We finally discuss how simulation models including trait variability within meta-PFTs are necessary to understand ecosystem responses to environmental drivers, both locally and globally and how this perspective will help to extend conceptual frameworks of other ecosystems to savanna research.     

Simulating Growth and Root-shoot Partitioning in Prairie Grasses Under Elevated Atmospheric CO2and Water Stress

We constructed a model simulating growth, shoot-root partitioning,plant nitrogen (N) concentration and total non-structural carbohydratesin perennial grasses. Carbon (C) allocation was based on theconcept of a functional balance between root and shoot growth,which responded to variable plant C and N supplies. Interactionsbetween the plant and environment were made explicit by wayof variables for soil water and soil inorganic N. The modelwas fitted to data on the growth of two species of perennialgrass subjected to elevated atmospheric CO₂and water stresstreatments. The model exhibited complex feedbacks between plantand environment, and the indirect effects of CO₂and water treatmentson soil water and soil inorganic N supplies were important ininterpreting observed plant responses. Growth was surprisinglyinsensitive to shoot-root partitioning in the model, apparentlybecause of the limited soil N supply, which weakened the expectedpositive relationship between root growth and total N uptake.Alternative models for the regulation of allocation betweenshoots and roots were objectively compared by using optimizationto find the least squares fit of each model to the data. Regulationby various combinations of C and N uptake rates, C and N substrateconcentrations, and shoot and root biomass gave nearly equivalentfits to the data, apparently because these variables were correlatedwith each other. A partitioning function that maximized growthpredicted too high a root to shoot ratio, suggesting that partitioningdid not serve to maximize growth under the conditions of theexperiment.Copyright 1998 Annals of Botany Company plant growth model, optimization, nitrogen, non-structural carbohydrates, carbon partitioning, elevated CO₂, water stress,Pascopyrum smithii,Bouteloua gracilis, photosynthetic pathway, maximal growth     

Cereal root exudates contain highly structurally complex polysaccharides with soil‐binding properties

Rhizosheaths function in plant?soil interactions, and are proposed to form due to a mix of soil particle entanglement in root hairs and the action of adhesive root exudates. The soil‐binding factors released into rhizospheres to form rhizosheaths have not been characterised. Analysis of the high‐molecular‐weight (HMW) root exudates of both wheat and maize plants indicate the presence of complex, highly branched polysaccharide components with a wide range of galactosyl, glucosyl and mannosyl linkages that do not directly reflect cereal root cell wall polysaccharide structures. Periodate oxidation indicates that it is the carbohydrate components of the HMW exudates that have soil‐binding properties. The root exudates contain xyloglucan (LM25), heteroxylan (LM11/LM27) and arabinogalactan‐protein (LM2) epitopes, and sandwich‐ELISA evidence indicates that, in wheat particularly, these can be interlinked in multi‐polysaccharide complexes. Using wheat as a model, exudate‐binding monoclonal antibodies have enabled the tracking of polysaccharide release along root axes of young seedlings, and their presence at root hair surfaces and in rhizosheaths. The observations indicate that specific root exudate polysaccharides, distinct from cell wall polysaccharides, are adhesive factors secreted by root axes, and that they contribute to the formation and stabilisation of cereal rhizosheaths.     

Spatial and temporal soil moisture resource partitioning by trees and grasses in a temperate savanna, Arizona, USA

Stable isotope analysis was used to determine sources of water used by coexisting trees and grasses in a temperate savanna dominated by Quercus emoryi Torr. We predicted that (1) tree seedlings and bunchgrasses utilize shallow sources of soil water, (2) mature savanna trees use deeper sources of water, and (3) trees switch from shallow to deep water sources within 1 year of germination. We found that Q. emoryi trees, saplings, and seedlings (about 2 months, 1 year, and 2 years old), and the dominant bunchgrass [Trachypogon montufari (H.B.K.) Nees.] utilized seasonally available moisture from different depths within the soil profile depending on size/age relationships. Sapling and mature Q. emoryi acquired water from >50 cm deep, 2-month-old seedlings utilized water from <15 cm, and 1- and 2-year-old seedlings and grasses used water from between 20 cm and 35 cm. This suggests that very young seedlings are decoupled from grasses in this system, which may facilitate germination and early establishment of Q. emoryi within extant stands of native grasses. The potential for subsequent interaction between Q. emoryi and native grasses was evidenced by similar patterns of soil water use by 1- and 2-year-old seedlings and grasses. Q. emoryi seedlings did not switch from shallow to deep sources of soil water within 2 years of germination: water use by these seedlings apparently becomes independent of water use by grasses after 2 years of age. Finally, older trees (saplings, mature trees) use water from deeper soil layers than grasses, which may facilitate the stable coexistence of mature trees and grasses. Potential shifts in the seasonality of precipitation may alter interactions between woody plants and grasses within temperate savannas characterized by bimodal precipitation regimes: reductions in summer precipitation or soil moisture may be particularly detrimental to warm-season grasses and seedlings of Q. emoryi. Received: 21 November 1996 / Accepted: 2 May 1997     

Water Uptake by Different Regions of the Barley Root. Pathways of Radial Flow in Relation to Development of the Endodermis

The water uptake by different lengths of lateral roots and 1.0cm or 5.0 cm lengths of the seminal axes from differentregionsof the root were measured in potometers with the shoot in airat two humidity regimes. A model of the contribution by thesedifferent regions of the root to the total water absorptionby the plant agreed well with measurements of water uptake bythe whole root system. According to this model, about one halfof the water taken up by the main axis came from the older suberizedregions further than 10 cm from the tip, and together with itsassociated lateral roots this region provided 75% of the totalwater transpired. The development of State III endodermal cellswas correlated with decreases in both the water uptake by theolder regions of the root and the translocation of calcium.Thus in the younger regions of the root where water uptake ismaximal, the flow of water is principally apoplastic althoughthere is also likely to be flow via the symplast Despite a 43%difference in transpirational demand between the two humiditytreatments, the leaf water potentials remained constant, implyinga change in root resistance. This change in resistance mightbe explained if there were an apoplastic pathway within thesuberin lamellae of State III endodermal cells. The responseto the increased transpirational demand is met by the olderregions of the root, in particular by the zone of lateral emergencewhere an apoplastic pathway is known to exist as the Casparianband in the endodermis breaks down with the emergence of thelateral roots. Key words: Endodermis, Pathways, Water Uptake