In situ measurement of fine root water absorption in three temperate tree species—Temporal variability and control by soil and atmospheric factors

Miniature heat balance-sap flow gauges were used to measure water flows in small-diameter roots (3–4 mm) in the undisturbed soil of a mature beech–oak–spruce mixed stand. By relating sap flow to the surface area of all branch fine roots distal to the gauge, we were able to calculate real time water uptake rates per root surface area (J_s) for individual fine root systems of 0.5–1.0 m in length. Study aims were (i) to quantify root water uptake of mature trees under field conditions with respect to average rates, and diurnal and seasonal changes of J_s, and (ii) to investigate the relationship between uptake and soil moisture θ, atmospheric saturation deficit D, and radiation I. On most days, water uptake followed the diurnal course of D with a mid-day peak and low night flow. Neighbouring roots of the same species differed up to 10-fold in their daily totals of J_s (<100–2000 g m⁻² d⁻¹) indicating a large spatial heterogeneity in uptake. Beech, oak and spruce roots revealed different seasonal patterns of water uptake although they were extracting water from the same soil volume. Multiple regression analyses on the influence of D, I and θ on root water uptake showed that D was the single most influential environmental factor in beech and oak (variable selection in 77% and 79% of the investigated roots), whereas D was less important in spruce roots (50% variable selection). A comparison of root water uptake with synchronous leaf transpiration (porometer data) indicated that average water fluxes per surface area in the beech and oak trees were about 2.5 and 5.5 times smaller on the uptake side (roots) than on the loss side (leaves) given that all branch roots <2 mm were equally participating in uptake. Beech fine roots showed maximal uptake rates on mid-summer days in the range of 48–205 g m⁻² h⁻¹ (i.e. 0.7–3.2 mmol m⁻² s⁻¹), oak of 12–160 g m⁻² h⁻¹ (0.2–2.5 mmol m⁻² s⁻¹). Maximal transpiration rates ranged from 3 to 5 and from 5 to 6 mmol m⁻² s⁻¹ for sun canopy leaves of beech and oak, respectively. We conclude that instantaneous rates of root water uptake in beech, oak and spruce trees are above all controlled by atmospheric factors. The effects of different root conductivities, soil moisture, and soil hydraulic properties become increasingly important if time spans longer than a week are considered.     

Reverse flow of sap in tree roots and downward siphoning of water by Grevillea robusta

1. Constant-power heat-balance sap flow gauges were used to compare sap flow in vertical and lateral roots of Grevillea robusta trees growing without access to ground water at a semiarid site in Kenya.
2. Reversal of sap flow occurred when root systems crossed gradients in soil water potential. Measurement of changes in the direction of flow was possible because of the symmetrical construction of the sap flow gauges; gradients in temperature across the gauges, and thus computed rates of sap flow, changed sign when reverse flow occurred.
3. Reverse flow in roots descending vertically from the base of the tree occurred, while uptake by lateral roots continued, when the top of the soil profile was wetter than the subsoil. The transfer of water downwards by root systems, from high to low soil water potential, was termed 'downward siphoning'; this is the reverse of hydraulic lift.
4. Downward siphoning was induced by the first rain at the end of the dry season and by irrigation of the soil surface during a dry period.
5. Downward siphoning may be an important component of the soil water balance where there are large gradients in water potential across root systems, from a wet soil surface downwards. By transferring water beyond the reach of shallow-rooted neighbours, downward siphoning may enhance the competitiveness of deep-rooted perennials.     

The role of soil hydraulic conductivity on the spatial and temporal variation of root water uptake in drip-irrigated corn

A multiplexed TDR system and a heat-pulse system for stem sap flow measurements were used to determine the spatial and temporal pattern of root water uptake in field-grown corn. The TDR probes, 0.15 and 0.30 m in length, were buried vertically in the soil profile to a depth of 0.95 m below the soil surface and heat-pulse sensors were installed on the plant base. Nocturnal readings from TDR probes were used successfully to differentiate the two components of moisture change: root uptake and net drainage. The instantaneous rate of water extraction by the plant measured by the heat-pulse system agreed well with the integrated rate of root water uptake measured frequently (at half-hour or hourly intervals) by the TDR probes. This agreement enabled further exploration into the cause of the evolution of the spatial and temporal patterns of root water uptake during a drying cycle. The results indicated that right after irrigation in the well-watered soil profile, it is the spatial distribution of the roots that mainly determines the typical pattern of root extraction, in addition to the fact that the roots near the plant base are more effective than those farther away. The higher density and effectiveness of the roots near the plant base dry the soil rapidly so that soil hydraulic conductivity soon becomes a limiting factor for water uptake. Further analysis revealed that a decrease in root uptake occurs near the plant base under a given atmospheric demand when the relative bulk soil hydraulic conductivity decreases to 0.002K _r. This suggests that low conductivity (high resistance) in the soil near the plant base is the initial cause for downward and lateral shifting of the root uptake pattern. Note that this critical value of hydraulic conductivity is not universal since it depends on the soil type and atmospheric water demand during the period under observation. Therefore, prior to the application of moisture content or suction head as measures of water availability or to control irrigation scheduling, it is suggested that these parameters be calibrated by the soil K() or K() curves, respectively, for the expected atmospheric water demand for the specific crop and growing period.     

A hydrological tracer study of water uptake depth in a Scots pine forest under two different water regimes

Little is known about the vertical distribution of water uptake by trees under different water supply regimes, the subject of this study, conducted in a Scots pine stand on sandy loam in northern Sweden. The objective was to determine the water uptake distribution in pines under two different water regimes, desiccation (no precipitation) and irrigation (2?mm day^?1 in July and 1?mm day^?1 in August), and to relate the uptake to water content, root and soil texture distributions. The natural ¹⁸O gradient in soil water was exploited, in combination with two added tracers, ²H at 10?cm and ³H at 20?cm depth. Extraction of xylem sap and water from the soil profile then enabled evaluation of relative water uptake from four different soil depths (humus layer, 0–10, 10–25 and 25–55?cm) in each of two 50-m² plots per treatment. In addition, water content, root biomass and soil texture were determined. There were differences in vertical water uptake distribution between treatments. In July, the pines at the irrigated and desiccated plots took up 50% and 30%, respectively, of their water from the upper layers, down to 25?cm depth. In August, the pines on the irrigated plots took up a greater proportion of their water from layers below 25?cm deep than they did in July. In a linear regression, the mean hydraulic conductivity for each mineral soil horizon explained a large part of the variation in relative water uptake. No systematic variation in the residual water uptake correlated to the root distribution. It was therefore concluded that the distribution of water uptake by the pines at Åheden was not a function of root density in the mineral soil, but was largely determined by the unsaturated hydraulic conductivity.     

Diurnal and seasonal variation in root xylem embolism in neotropical savanna woody species: impact on stomatal control of plant water status

Vulnerability to water-stress-induced embolism and variation in the degree of native embolism were measured in lateral roots of four co-occurring neotropical savanna tree species. Root embolism varied diurnally and seasonally. Late in the dry season, loss of root xylem conductivity reached 80% in the afternoon when root water potential (psi root) was about -2.6 MPa, and recovered to 25-40% loss of conductivity in the morning when psi root was about -1.0 MPa. Daily variation in psi root decreased, and root xylem vulnerability and capacitance increased with rooting depth. However, all species experienced seasonal minimum psi root close to complete hydraulic failure independent of their rooting depth or resistance to embolism. Predawn psi root was lower than psi soil when psi soil was relatively high (> -0.7 MPa) but became less negative than psi soil, later in the dry season, consistent with a transition from a disequilibrium between plant and soil psi induced by nocturnal transpiration to one induced by hydraulic redistribution of water from deeper soil layers. Shallow longitudinal root incisions external to the xylem prevented reversal of embolism overnight, suggesting that root mechanical integrity was necessary for recovery, consistent with the hypothesis that if embolism is a function of tension, refilling may be a function of internal pressure imbalances. All species shared a common relationship in which maximum daily stomatal conductance declined linearly with increasing afternoon loss of root conductivity over the course of the dry season. Daily embolism and refilling in roots is a common occurrence and thus may be an inherent component of a hydraulic signaling mechanism enabling stomata to maintain the integrity of the hydraulic pipeline in long-lived structures such as stems.     

Root pressure and specific conductivity in temperate lianas: exotic Celastrus orbiculatus (Celastraceae) vs. native Vitis riparia (Vitaceae)

The exotic temperate liana (woody vine) Celastrus orbiculatus has become a weed in Michigan, occurring in many of the same habitats as the native liana Vitis riparia. However, C. orbiculatus frequently develops into extensive monospecific infestations, while V. riparia does not. Freezing-induced embolism may be responsible for limiting liana distribution. Root pressure has been observed in numerous tropical lianas and temperate species of Vitis and has been implicated as vital to the recovery of xylem function in wide vessels following winter freezes. For both of these co-occurring lianas we investigated root pressure and water conductance as possible explanatory factors for their differential spread. According to our hypothesis, C. orbiculatus should have produced greater or more frequent root pressures than V. riparia. However, the reverse proved true, indicating that root pressure is not a prerequisite for weedy proliferation of C. orbiculatus. Additionally, the seasonal patterns of specific conductivity of stem xylem indicate that each species responds differently to environmental constraints. Vitis riparia establishes conductivity early in the growing season, before the leaves emerge, using root pressure to reverse embolism, but loses conductivity with the first freeze in early autumn. Celastrus orbiculatus is slow to establish conductivity, depending on new wood production, but leafs out sooner than V. riparia and maintains green leaves after the first freeze. Vulnerability curves of xylem to cavitation caused by water stress for the two species indicate that they respond similarly to dehydration. These results indicate that root pressures are not responsible for the invasive success of C. orbiculatus and suggest that other factors must be key to its prolific invasion.     

Hydraulic redistribution of soil water in two old-growth coniferous forests: quantifying patterns and controls

Although hydraulic redistribution of soil water (HR) by roots is a widespread phenomenon, the processes governing spatial and temporal patterns of HR are not well understood. We incorporated soil/plant biophysical properties into a simple model based on Darcy's law to predict seasonal trajectories of HR. We investigated the spatial and temporal variability of HR across multiple years in two old-growth coniferous forest ecosystems with contrasting species and moisture regimes by measurement of soil water content (theta) and water potential (Psi) throughout the upper soil profile, root distribution and conductivity, and relevant climate variables. Large HR variability within sites (0-0.5 mm d(-1)) was attributed to spatial patterns of roots, soil moisture and depletion. HR accounted for 3-9% of estimated total site water depletion seasonally, peaking at 0.16 mm d(-1) (ponderosa pine; Pinus ponderosa) or 0.30 mm d(-1) (Douglas-fir; Pseudotsuga menziesii), then declining as modeled pathway conductance dropped with increasing root cavitation. While HR can vary tremendously within a site, among years and among ecosystems, this variability can be explained by natural variability in Psi gradients and seasonal courses of root conductivity.     

Secondary root growth in Rhynchosia edulis Griseb. (Leguminosae): Origin of cambia and their products

This study focuses on the development of a secondary root structure in Rhynchosia edulis Griseb. (Leguminosae). Its principal objectives are (i) to study the origin of cambia and the nature of their products; (ii) to correlate root structure to habitat; and (iii) to compare this anatomy with that of other Leguminosae species growing in the same environment. Serial transverse cuts of the main root show that the secondary root structure in this species results from several phenomena, namely (1) a cambium arising from procambial and pericycle cells; (2) a lateral meristem producing cell layers from the periphery towards the inner part of the root and from which vascular bundles, whose cambia fuse forming a continuous ring, originate; and (3) the formation of “elliptical cambia” in the mid portion of the root giving rise to vascular bundles in reverse orientation. The comparison of secondary root growth in R. edulis with other root structures in Leguminosae species growing in hilly areas shows different structural patterns. Nonetheless, these different patterns have the same objective: to enlarge storage parenchyma tissue enabling survival within an environment having limited water availability.     

MRI of long-distance water transport: a comparison of the phloem and xylem flow characteristics and dynamics in poplar, castor bean, tomato and tobacco

We used dedicated magnetic resonance imaging (MRI) equipment and methods to study phloem and xylem transport in large potted plants. Quantitative flow profiles were obtained on a per-pixel basis, giving parameter maps of velocity, flow-conducting area and volume flow (flux). The diurnal xylem and phloem flow dynamics in poplar, castor bean, tomato and tobacco were compared. In poplar, clear diurnal differences in phloem flow profile were found, but phloem flux remained constant. In tomato, only small diurnal differences in flow profile were observed. In castor bean and tobacco, phloem flow remained unchanged. In all plants, xylem flow profiles showed large diurnal variation. Decreases in xylem flux were accompanied by a decrease in velocity and flow-conducting area. The diurnal changes in flow-conducting area of phloem and xylem could not be explained by pressure-dependent elastic changes in conduit diameter. The phloem to xylem flux ratio reflects what fraction of xylem water is used for phloem transport (Münch's counterflow). This ratio was large at night for poplar (0.19), castor bean (0.37) and tobacco (0.55), but low in tomato (0.04). The differences in phloem flow velocity between the four species, as well as within a diurnal cycle, were remarkably small (0.25-0.40 mm s(-1)). We hypothesize that upper and lower bounds for phloem flow velocity may exist: when phloem flow velocity is too high, parietal organelles may be stripped away from sieve tube walls; when sap flow is too slow or is highly variable, phloem-borne signalling could become unpredictable.     

Resistance to water flow in xylem of Picea abies (L.) Karst. trees grown under contrasting light conditions

Summary The relative hydraulic conductivity (k) of xylem and resistance (R) to water flow through trunk, primary roots and branches in Picea abies trees growing under contrasting light conditions were investigated. The xylem permeability to water was measured by forcing 10 mM water solution of KC1 through excised wood specimens. From the values of k, the sapwood transverse area and the length of conducting segments, R of the whole trunk, branches and roots was calculated. The relative conductivity of xylem in open-grown trees exceeded that of shade-grown trees by 1.4–3.1 times, while k was closely correlated with the hydraulically effective radius (R _e) of the largest tracheids (R ² was 0.85–0.94 for open- and 0.51–0.79 for shade-grown trees). Because of both a low k and a smaller sapwood area in shade-grown trees the resistance to water movement through their trunk, roots and branches was many times higher. The distribution of R between single segments of the water-conducting pathway differed considerably in trees from different sites. At high water status the largest share of the total resistance in open- as well as shade-grown trees resides in the apical part of the trunk. The contribution of the branches to total xylem resistance is supposed to increase with developing water deficit.     

Conservation of two predator species for biological control ofChrysophtharta bimaculata (Col.: Chrysomelidae) in tasmanian forests

Chrysophtharta bimaculata (Olivier) (Coleoptera: Chrysomelidae) is a major defoliator of regeneration eucalypt trees in Tasmania causing a significant reduction in height and diameter increment of trees which reduces wood volume per hectare. A study to conserve and enhance the efficiency of coccinellid species chieflyCleobora mellyi (Mulsant) (Coleoptera: Coccinellidae), and the cantharid,Chauliognathus pulchellus (Macleay) (Coleoptera: Cantharidae), for the biological control ofC. bimaculata was conducted in young regeneration forests in southern Tasmania from 1991–92. Cantharid adults and coccinellid adults and larvae feed onC. bimaculata eggs and, to a lesser extent, young larvae. The study found that coccinellids were more active throughout the egg and early (1^st and 2^nd) stage ofC. bimaculata. The cantharid, however was active only during the egg stage of the prey and then disappeared from the plantation. The coccinellids were therefore the most useful predators, but their population declined when the prey reached the 3^rd and 4^th stages. As shortage of food may account for this decline, supplementary food was provided in the form of sucrose sprays or sugar granules at a feeding station. This resulted in the retention of both predators and particularly the coccinellids and enhanced their efficacy.     

Importation,releases, and establishment ofNeochetina spp. (Col.: Curculionidae) for the biological control of water hyacinth,Ecchhornia crassipes (Lil.: Pontederiaceae), in Benin,West Africa

Water hyacinth,Eichhornia crassipes (Martius) Solms-Laubach (Pontederiaceae) was first reported in Bénin in 1977 and about 10 years later became the major floating water weed in the south east, obstructing boat traffic and fisheries. Water hyacinth multiplies in permanently fresh water in the swampy upper reaches of the Sô River and in tributaries of the Ouémé River. From there it is moved by wind and water flow to the coastal lagoons. The coastal lagoons are brackish during the dry season and water hyacinth eventually dies. In 1991,Neochetina eichhorniae (Warner) (Col.: Curculionidae) of South American origin was imported from Australia via quarantine in Britain to Bénin. A small infestation of the fungusBeauveria bassiana (Bals.) Vuill. (Hyphomycetes) was eliminated from the colony before release by sterilizing eggs and rearing a fungus-free generation. Between late 1991 and mid 1993, about 23,900N. eichhorniae were released at 11 localities along the Ouémé River and in the head waters of the Sô River. Regular monitoring revealed feeding scars by adults on leaves and tunnelling by larvae in petioles at all release sites. By October 1993,N. eichhorniae had spread up to 20 km from some release sites.Neochetina bruchi Hustache was imported in 1992. A total of about 5,700 weevils has been released in six localities since mid 1992. Recoveries of offspring were made in all but one locality. Despite the negative impact of water flow, wind, penetration of salt water, and removal of infested water hyacinth by fishermen,N. eichhorniae andN. bruchi are established in Bénin in a situation typical for coastal West Africa.