Influences of Water Status, Temperature, and Root Age on Daily Patterns of Root Respiration for Two Cactus Species

Daily patterns of root respiration measured as CO₂, efflux werestudied at various soil water potentials, temperatures, androot ages for individual, attached roots of the barrel cactusFerocactus acanthodes and the platyopuntia Opuntia ficus-indica.The daily patterns of root respiration for both establishedroots and rain roots followed the daily patterns of root temperature.Root respiration increased when root temperature was raisedfrom 5 °C to 50 °C for F. acanthodes and from 5 °Cto 55 °C for O. ficus-indica; at 60 °C root respirationdecreased 50° from the maximum for F. acanthodes and decreased25° for O. ficus-indica. Root respiration per unit d. wtdecreased with root age for both species, especially for rainroots. Root respiration rates for rain roots were reduced tozero at a soil water potential (     

Water Relations and Photosynthesis of a Desert CAM Plant, Agave deserti

The water relations and photosynthesis of Agave deserti Engelm., a plant exhibiting Crassulacean acid metabolism, were measured in the Colorado desert. Although no natural stomatal opening of A. deserti occurred in the summer of 1975, it could be induced by watering. The resistance for water vapor diffusion from a leaf (R_WV) became less than 20 sec cm⁻¹ when the soil water potential at 10 cm became greater than −3 bars, as would occur after a 7-mm rainfall. As a consequence of its shallow root system (mean depth of 8 cm), A. deserti responded rapidly to the infrequent rains, and the succulent nature of its leaves allowed stomatal opening to continue for up to 8 days after the soil became drier than the plant. When the leaf temperature at night was increased from 5 to 20 C, R_WV increased 5-fold, emphasizing the importance of cool nighttime temperatures for gas exchange by this plant. Although most CO₂ uptake occurred at night, a secondary light-dependent rise in CO₂ influx generally occurred after dawn. The transpiration ratio (mass of water transpired/mass of CO₂ fixed) had extremely low values of 18 for a winter day, and approximately 25 for an entire year.     

Root Hydraulic Conductivity of Two Cactus Species in Relation to Root Age, Temperature, and Soil Water Status

The effects of root age, temperature, and soil water statuson root hydraulic conductivity (L_P) were investigated for twocactus species, Ferocactus acanthodes and Opuntia ficus-indica.The volumetric flux density of water was measured for excisedroot segments, either using negative hydrostatic pressures appliedto the proximal end or using reverse flow of water from theroot to the soil. For both species, L_P at 20 ?C increased withroot age, average values reaching a maximum of 3.9 ? 10^–7m s^–1 MPa^–1 for F. acanthodes and 5.2 ? 10^–7m s^–1 MPa^–1 for O.ficus-indica at 11 to 17 weeksof age; L_P subsequently declined with increasing root age forboth species. L_P was maximal at a temperature of about 10 ?Cfor the youngest roots (1–3 weeks), this optimum shiftingto 40 ?C for 8-week-old roots of both species. For older roots(up to 1.5-years-old), L_P increased with temperature from 0?C to 50 ?C, with a Q₁₀ of 1.3 between 20 ?C and 30 ?C. At asoil water potential (_soil) of –0.016 MPa, root L_P wasindependent of the direction of water flow for both species.Depending on root age, L_P declined 45- to 500-fold for F. acanthodesand 90- to 800-fold for O.ficus-indica as _soil was reduced from–0.016 to –1.06 MPa, consistent with a rectifier-likebehaviour with respect to water movement between soil and roots.Incorporation of such responses into water uptake models shouldlead to a better understanding of root function. Key words: Ferocactus acanthodes, Opuntia ficus-indica, water potential, tension, reverse flow     

Water Influx Characteristics and Hydraulic Conductivity for Roots of Agave deserti Engelm.

Various plant and environmental factors influence the hydraulicproperties for roots, which were examined using negative hydrostaticpressures applied to the proximal ends of individual excisedroots of a common succulent perennial from the Sonoran Desert,Agave deserti Engelm. The root hydraulic conductivity, L^p, increasedsubstantially with temperature, the approximately 4-fold increasefrom 0.5°C to 40°C representing a Q¹⁰ of 1.45. Suchvariations in L_p with temperature must be taken into accountwhen modelling water uptake, as soil temperatures in the rootzone of such a shallow-rooted species vary substantially bothdaily and seasonally. At 20°C, L_p was 2.3 x 10^–7 ms^{macron}1MPa^{macron}1for 3-week-old roots, decreasing to abouthalf this value at 10 weeks and then becoming approximatelyhalved again at 6 months. For a given root age, L_p for rainroots that are induced by watering as lateral branches on theestablished roots (which arise from the stem base) was aboutthe same as L_p for established roots. Hence, the conventionalbelief that rain roots have a higher L_p than do establishedroots is more a reflection of root age, as the rain roots tendto be shed following drought and thus on average are much youngerthan are established roots. Unlike previous measurements onroot respiration, lowering the gas-phase oxygen concentrationfrom 21% to 0% or raising the carbon dioxide concentration from0.1% to 2% had no detectable effect on L_p for rain roots andestablished roots. L_p for rain roots and established roots wasdecreased by an average of 11% and 35% by lowering the soilwater potential from wet conditions (_soil=0 kPa) to {macron}40kPa and {macron}80 kPa, respectively. Such decreases in L_p mayreflect reduced water contact between soil particles and theroot surface and should be taken into account when predictingwater uptake by A. deserti. Key words: Gas phase, rain roots, root age, soil, temperature, water potential     

Hydraulic Conductivity and Anatomy for Lateral Roots of Agave deserti During Root Growth and Drought-induced Abscission

Hydraulic conductivity (L_p), radial conductivity (L_R), axialconductance (K_h), and related anatomical characteristics forlateral roots of Agave deserti were investigated during rootgrowth and drought-induced abscission. The elongation rate oflateral roots averaged 5 mm d^–1 under wet conditions andwas reduced 95% by 17 d of drought (     

Temperature,water, and PAR influences on predicted and measured productivity of Agave deserti at various elevations

Summary To measure productivity of Agave deserti over its elevational range in the northwestern Sonoran Desert, leaf unfolding from its basal rosette was monitored on groups of 10 plants at 13 sites. Based on data from an intermediate elevation (840 m), leaf unfolding proved to be highly correlated (r ²=0.88) with an environmental productivity index (EPI) formed as the product of indices for water status, temperature, and photosynthetically active radiation (PAR); each of these latter indices indicated the fraction of maximum net CO₂ uptake expected for that parameter based on laboratory measurements of gas exchange and field microclimatic data. At 840 m, the main environmental variable influencing leaf unfolding for A. deserti over a 2-y period was soil water potential. On steep slopes, however, leaf unfolding during the winter ranged from 0.7 leaves per 10 plants for north-facing slopes to 7.3 for south-facing slopes, reflecting the importance of PAR. Summer and winter rainfall increased 3-fold from elevations of 300 m to 1,200 m. Temperatures were more optimal for net CO₂ uptake at high elevations in the summer and at low elevations in the winter. Hence EPI increased with elevation in the summer but was maximal at an intermediate elevation in the winter. Moreover, measured leaf unfolding in both the summer and the winter closely followed the changes in EPI with elevation, indicating that productivity could be closely predicted for A. deserti based on physiological CO₂ responses and changes in environmental conditions with elevation.     

Water Movement and Storage in a Desert Succulent: Anatomy and Rehydration Kinetics for Leaves of Agave deserti

Smith, J. A. C. and Nobel, P. S. 1986. Water movement and storagein a desert succulent: anatomy and rehydration kinetics forleaves of Agave deserti.—J. exp. Bot. 37: 1044–1053. Anatomic and kinetic aspects of water storage were investigatedfor the succulent leaves of the desert CAM plant, Agave deserti.An approximately linear relationship was found between the numberof vascular bundles and leaf surface area, both for leaves ofdifferent sizes and also along the length of a single leaf.The bundles, which were distributed throughout the leaf cross-section,were separated from each other by about eight water-storagecells. Even though the cell walls of the water-storage groundtissue made up only 2?5% of the cell volume, they provided about10% of the total cross-sectional area available for water transportradial to the xylem because cell-cell contact in such a directionaveraged 25% of the cell surface area. The rehydration kineticsof partially dehydrated leaf segments were resolved into threephases: (1) a relatively rapid movement into the vascular tissue(half-time of 2 min); (2) water movement into storage in theground tissue (half-time of 59 min); and (3) water movementinto the intercellular air spaces (half-time of about 10 h).Using the observed kinetics for water movement into the storagetissue and standard diffusion theory, the bulk-averaged diffusivityof water in the relatively homogeneous ground tissue (D₁) was2?0 ? 10^–10 m² s^–1 Using this (D₁) and pathway analysis,most of the water moving from the xylem into storage in themassive leaves of A. deserti apparently occurred from cell tocell across the cell membranes rather than through the cellwalls. Key words: Agave deserti, capacitance, diffusivity, leaf anatomy, succulence, water storage     

Water flow and water storage in Agave deserti: osmotic implications of crassulacean acid metabolism

Abstract Water flow and water storage were investigated for Agave deserti, a desert succulent showing crassulacean acid metabolism (CAM). The anatomy and water relations of the peripheral chlorenchyma, where CAM occurs, and the central water-storage parenchyma were investigated for its massive leaves so that these tissues could be incorporated as discrete elements into an electrical-circuit analogue of the whole plant. The daily cycling of osmotic pressure was represented by voltage sources in series with the storage capacitors. With soil water potential and leaf transpiration rate as input variables, axial water flow through the vascular bundles and radial flows into and out of storage during the day/night cycle were determined. The predominantly nocturnal transpiration was coincident with increases in cell osmotic pressure and in titratable acid of the leaf chlorenchyma. In the outer layers of the chlorenchyma, water potential was most negative at the beginning of the night when transpiration was maximum, while the water-storage parenchyma reached its minimal water potential 9 h later. The roots plus stem contributed 7% and the leaves contributed 50% to the total water flow during maximal transpiration; peak water flow from the soil to the roots occurred at dawn and was only 58% of the maximal transpiration rate. Over each 24-h period, 39% of the water lost from the plant was derived from storage, with flow into storage occurring mainly during the daytime. Simulations showed that the acid accumulation rhythm of CAM had little impact on water uptake from the soil under the conditions employed. In the outer chlorenchyma, water potential and water flows were more sensitive to the day/night changes in transpiration than in osmotic pressure. Nevertheless, cell osmotic pressure had a large influence on turgor pressure in this tissue and determined the extent to which storage was recharged during the latter part of the night.     

Parent-ramet connections in Agave deserti: influences of carbohydrates on growth

Summary Agave deserti, a monocarpic perennial occurring in the northwestern Sonoran Desert, produces ramets on rhizomes that extend from the base of a parent plant. Shading ramtes to light compensation for two years did not decrease their relative growth rate (RGR) compared with unshaded ramets. However, the parents experienced a 30% decrease in total nonstructural carbohydrate (TNC) level, suggesting that carbohydrates were translocated from parents to ramets. Shaded parents had RGR's similar to unshaded parents, due in large part to consumption of 50% of the TNC reserves of shaded parents, but about 10% of the growth of the shaded parents was attributed to TNC received from their attached ramets. Estimates of parent and ramet growth separately, based on changes in TNC levels (converted to dry weight using a measured production value), net CO₂ uptake of unfolded leaves, and respiration of roots, stems, and folded leaves, were similar to measured growth of parents and ramets combined, suggesting that parents and ramets are physiologically integrated and grow as a unit. Large TNC reserves, which were also shown to support growth during conditions of water limitation in the field, enhance the growth of ramets in an environment where seedling establishment is rare.Abbreviations DW dry weight - EPI environmental productivity index - PAR photosynthetically active radiation - RGR relative growth rate - TNC total nonstructural carbohydrate     

Influence of Daily Short-Term Temperature Drops on Respiration to Photosynthesis Ratio in Chilling-Sensitive Plants

Cucumber (Cucumis sativus L.), tomato (Solanum lycopersicum L.), and sweet pepper (Capsicum annuum L.) plants were subjected daily over 13 days to short-term (2 h) temperature drops to 12, 8, 4, and 1°C (DROP treatments) at the end of night periods, and effects of these chilling treatments on the ratio of dark respiration in leaves (R_d) to gross photosynthesis (A_g) were examined. The results showed that DROP treatments affected the R_d/A_g ratio in leaves: this ratio increased significantly in cucumber and tomato plants and was slightly affected in pepper plants. When the temperature drops to 12°C were applied, the increase in R_d/A_g ratio in cucumber and tomato plants was entirely due to the rise in R_d. In the case of temperature drops to 8°C and below, the increase in R_d/A_g was determined by both elevation of R_d and the concurrent decrease in Ag. In cucumber plants, the extent of Ag and R_d changes increased with the DROP severity, i.e., with lowering the temperature of DROP treatment. The inhibition of photosynthesis by DROP treatment in cucumber plants was accompanied by the diminished efficiency of light energy use for photosynthesis and by the increase in the light compensation point. The elevation in R_d/A_g ratio in cucumber plants was accompanied by the decline in growth characteristics, such as accumulation of aboveground biomass, plant height, and leaf area. It was concluded that the R/A ratio is an important indicator characterizing the adaptive potential of chilling-sensitive plant species and their response to daily short-term temperature drops.     

Environmental Influences on Open Stomates of a Crassulacean Acid Metabolism Plant, Agave deserti

The major short term stomatal response of Agave deserti was to temperature; increases in leaf temperature led to decreases in water vapor conductance for stomatal opening during the daytime (C₃ mode) as well as at night (Crassulacean acid metabolism or CAM mode). Hourly changes in the water vapor concentration drop from leaf to air had no significant stomatal effect in either mode. Stomatal responses to external CO₂ levels up to 800 microliters per liter were not significant after 15 minutes and only moderate after a few hours, suggesting that CO₂ effects on open stomates of this succulent were indirect in both CAM and C₃ modes.     

Influence of soil O2 and CO2 on root respiration for Agave deserti

Respiration measured as CO₂ efflux was determined at various soil O₂ and CO₂ concentrations for individual, attached roots of a succulent perennial from the Sonoran Desert, Agave deserti Engelm. The respiration rate increased with increasing O₂ concentration up to about 16% O₂ for established roots and 5% O₂ for rain roots (fine branch roots on established roots induced by wetting of the soil) and then remained fairly constant up to 21% O₂. When O₂ was decreased from 21 to 0%, the respiration rates were similar to those obtained with increasing O₂ concentration. The CO₂ concentration in the root zone, which for the shallow-rooted A. deserti in the field was about 1 000 μl l^-1, did not affect root respiration at concentrations up to 2 000 μl l^-1, but higher concentrations reduced it, respiration being abolished at 20 000 μl l^-1 (2%) CO₂ for both established and rain roots. Upon lowering CO₂ to 1 000 μl l^-1 after exposure to concentrations up to 10000 μl l^-1 CO₂, inhibition of respiration was reversible. Uptake of the vital stain neutral red by root cortical cells was reduced to zero, indicating cell death, in about 4 h at 2% CO₂, substantiating the detrimental effects of high soil CO₂ concentrations on roots of A. deserti . This CO₂ response may explain why roots of desert succulents tend to occur in porous, well-aerated soils.     

Aquaporins account for variations in hydraulic conductance for metabolically active root regions of Agave deserti in wet, dry, and rewetted soil

The importance of aquaporins for root hydraulic conductance (L_P) was investigated along roots of the desert succulent Agave deserti in wet, dry and rewetted soil. Water channel activity was inferred from HgCl₂‐induced reductions of L_P that were reversible by 2‐mercaptoethanol. Under wet conditions, HgCl₂ reduced L_P for the distal root region by 50% and for the root region near the shoot base by 36% but did not affect L_P for the mid‐root region. For all root regions, L_P decreased by 30–60% during 10 d in drying soil and was not further reduced by HgCl₂. After soil rewetting, L_P increased to pre‐drying values and was again reduced by HgCl₂ for the distal and the basal root regions but not the mid‐root region. For the distal region, water channels in the epidermis/exodermis made a disproportionately large contribution to radial hydraulic conductance of the intact segment; for the basal region, water channel activity was highest in the cortex and endodermis. The role of water channels was greatest in tissues in which cells were metabolically active both in the distal root region, where new apical growth occurs in wet soil, and in the basal region, which is the most likely root region to intercept light rainfall.     

Productivity of Agave deserti: measurement by dry weight and monthly prediction using physiological responses to environmental parameters

Summary An environmental productivity index based on physiological responses to three environmental variables was used to predict the net productivity of a common succulent perennial of the Sonoran Desert, Agave deserti, on a monthly basis. Productivity was also independently measured in the field from dry weight changes. The index was based on soil water availability, day/night air temperatures, and photosynthetically active radiation (PAR), which were individually varied in the laboratory and the effect on net CO₂ uptake by the leaves determined. From monthly precipitation, temperature, and PAR at the field site together with the responses measured in the laboratory, an index (maximum value of unity) was assigned to each of these three environmental variables and their product was termed the environmental productivity index. This index indicates the fraction of maximal CO₂ uptake expected in the field for each month (well-watered A. deserti assimilated 285 mmol CO₂ m^-2 leaf area day^-1 at PAR saturation and optimal day/night temperatures of 25° C/15° C). The dry weight analysis was based on the monthly unfolding of new leaves from the central spike of the rosette and their seasonal increase in dry weight, which were determined in the field. The production of new leaves was highly correlated with the environmental productivity index (r²=0.93), which in turn was highly correlated with the water status index (r²=0.97). After correction for respiration by folded leaves, stem, and roots, plant productivity predicted by the average environmental productivity index (0.36) over a wet June-to-October period agreed within 4% with the productivity based on the conventional dry weight analysis. The net productivity of A. deserti over this 5-month period was 0.57 kg m^-2 ground area (5.7 Mg ha^-1), a large value for a desert CAM plant. The environmental productivity index proposed here may provide a reliable means for predicting net productivity on a monthly basis, which may be particularly useful for species in relatively variable environments such as deserts.     

羊草呼吸作用与湿度、光照和土壤水分的关系

本文报道了两种土壤水分条件下羊草明呼吸速率与光照和温度的关系,以及暗呼吸速率与温度的关系。结果表明,羊草的明呼吸速率与光强呈非线性函数关系。在低光强下,明呼吸速率随光强升高而有较快的增加;随着光强的增高,其增加速度减慢。在温度低于羊草光合的高温补偿点的条件下,明呼吸速率在一定温度范围内随温度升高而增大;当温度达到一定限度时,有一个下降阶段,而后又回升,羊草的暗呼吸速率随温度增加而升高,且在一定限度内,其升高速度随温度增高而加快。当土壤干旱时,明呼吸速率显著降低,而暗呼吸速率仅略有减小。     

Temperature influences on root growth for Encelia farinosa (Asteraceae), Pleuraphis rigida (Poaceae), and Agave deserti (Agavaceae) under current and doubled CO2 concentrations

To help evaluate root distribution patterns, elongation rates of individual roots were measured as a function of soil temperature for Encelia farinosa (a C₃ species), Pleuraphis rigida (C₄), and Agave deserti (CAM), sympatric codominants in the northwestern Sonoran Desert. Measurements were made at current and doubled CO₂ concentrations under winter and summer conditions of air temperature (day/night temperatures of 17 C/10 C and 33 C/22 C, respectively). The three species had different optimal temperatures for root elongation (T_opt) under winter conditions (25 C for E. farinosa, 35 C for P. rigida, and 30 C for A. deserti); T_opt increased by 2-3 C under summer conditions for all three species. The limiting temperatures for elongation also acclimated from winter to summer conditions. The rate of root elongation at T_opt was higher under summer than winter conditions for E. farinosa (9 vs. 6 mm d⁻¹) and P. rigida (20 vs. 14 mm d⁻¹), reflecting conditions for maximum photosynthesis; no difference occurred for A. deserti (9 vs. 10 mm d⁻¹). Decreased elongation rates at extreme temperatures were associated with less cell division and reduced cell extension. The doubled CO₂ concentration increased average daily root elongation rates for A. deserti under both winter (7%) and summer (12%) conditions, reflecting increased cell extension, but had no effect for the other two species. Simulations of root elongation as a function of soil temperatures showed that maximum elongation would occur at different depths (16-20 cm for E. farinosa, 4-8 cm for P. rigida, and 0-4 cm for A. deserti) and during different seasons (winter to spring for E. farinosa, spring to summer for P. rigida, and all year for A. deserti), contributing to their niche separation. Shading of the soil surface moderated daily variations in soil temperature, reducing seasonal root elongation for winter and spring and increasing elongation for summer. Shading also altered root distribution patterns, e.g., optimal rooting depth for A. deserti and especially P. rigida increased for a hot summer day.     

Root Respiration and Carbohydrate Status of Two Wheat Genotypes in Response to Hypoxia

To investigate root respiration and carbohydrate status in relationto waterlogging or hypoxia tolerance, root respiration rateand concentrations of soluble sugars in leaves and roots weredetermined for two wheat (Triticum aestivum L.) genotypes differingin waterlogging-tolerance under hypoxia (5% O₂) and subsequentresumption of full aeration. Root and shoot growth were reducedby hypoxia to a larger extent for waterlogging-sensitive Coker9835. Root respiration or oxygen consumption rate declined withhypoxia, but recovered after 7 d of resumption of aeration.Respiration rate was greater for sensitive Coker 9835 than fortolerant Jackson within 8 d after hypoxia. The concentrationsof sucrose, glucose and fructose decreased in leaves for bothgenotypes under hypoxia. The concentration of these sugars inroots, however, increased under hypoxia, to a greater degreefor Jackson. An increase in the ratio of root sugar concentrationto shoot sugar concentration was found for Jackson under hypoxicconditions, suggesting that a large amount of carbohydrate waspartitioned to roots under hypoxia. The results indicated thatroot carbohydrate supply was not a limiting factor for rootgrowth and respiration under hypoxia. Plant tolerance to waterloggingof hypoxia appeared to be associated with low root respirationor oxygen consumption rate and high sugar accumulation underhypoxic conditions.Copyright 1995, 1999 Academic Press Oxygen consumption rate, sugar accumulation, Triticum aestivum L., waterlogging tolerance