The decrease in growth of phosphorus-deficient maize leaves is related to a lower cell production

The spatial distribution of leaf elongation and adaxial epidermal cell production in leaf 6 of maize (Zea mays L. cv. Cecilia) plants grown in a growth chamber under two contrasting availabilities of P in the soil was investigated. Lower displacement velocities from 32.5 mm from leaf base and a shorter growth zone were found in low P (LP) leaves compared with control leaves. P deficiency significantly diminished maximum relative elemental growth rate and shifted its location closer to the leaf base. Cells were significantly longer in LP than in control leaves for all positions from the leaf base except at the end of the growth zone. For both treatments it took a similar time for a cell situated at the leaf base to reach the limit of the growth zone. The average length of the cell division zone was decreased by 21% in LP leaves. Significant differences were found in cell production and cell division rates from 12.5 mm from the leaf base although maximum values were similar between P treatments. A shorter zone of cell division with lower cell production rates along most of its length was the regulatory event that decreased cell production, and ultimately leaf elongation rates, in P‐deficient maize plants.     

Spatial distribution of growth rates and of epidermal cell lengths in the elongation zone during leaf development in Lolium perenne L.

Relative elemental growth rates (REGR) and lengths of epidermal cells along the elongation zone of Lolium perenne L. leaves were determined at four developmental stages ranging from shortly after emergence of the leaf tip to shortly before cessation of leaf growth. Plants were grown at constant light and temperature. At all developmental stages the length of epidermal cells in the elongation zone of both the blade and sheath increased from 12 m at the leaf base to about 550 m at the distal end of the elongation zone, whereas the length of epidermal cells within the joint region only increased from 12 to 40 m. Throughout the developmental stages elongation was confined to the basal 20 to 30 mm of the leaf with maximum REGR occurring near the center of the elongation zone. Leaf elongation rate (LER) and the spatial distributions of REGR and epidermal cell lengths were steady to a first approximation between emergence of the leaf tip and transition from blade to sheath growth. Elongation of epidermal cells in the sheath started immediately after the onset of elongation of the most proximal blade epidermal cells. During transition from blade to sheath growth the length of the blade and sheath portion of the elongation zone decreased and increased, respectively, with the total length of the elongation zone and the spatial distribution of REGR staying near constant, with exception of the joint region which elongated little during displacement through the elongation zone. Leaf elongation rate decreased rapidly during the phase when only the sheath was growing. This was associated with decreasing REGR and only a small decrease in the length of the elongation zone. Data on the spatial distributions of growth rates and of epidermal cell lengths during blade elongation were used to derive the temporal pattern of epidermal cell elongation. These data demonstrate that the elongation rate of an epidermal cell increased for days and that cessation of epidermal cell elongation was an abrupt event with cell elongation rate declining from maximum to zero within less than 10 h.Abbreviations LER leaf elongation rate - REGR relative elemental growth rates     

Effect of N deficiency on leaf growth and cell wall peroxidase activity in contrasting maize genotypes

Two maize genotypes differing in leaf elongation rate (high-LER and low-LER) were used for the investigation of the effects of nitrogen deficiency on leaf growth and development and activity of enzyme cell wall peroxidase in the leaf growth zone. Plants were grown in a growth cabinet in perlite as a substrate and watered with complete N-NO₃ solution (+N) and N-NO₃ deficient solution (–N). Comparison between the investigated genotypes showed that final leaf length in both N treatments was related with LER, but not with the duration of leaf elongation. Faster leaf elongation rate in high-LER compared with low-LER genotype, was associated with longer growth zone, a bigger number of cells in it, and higher cell flux rate, although cell elongation rate was similar in both genotypes. These lines of evidence indirectly indicated that leaves of the faster growing genotype were characterized by higher meristematic activity. Nitrogen deficiency reduced the flux of cells and cell elongation rate, length of cell division zone and the number of cells in whole zone, significantly for both genotypes, although duration of cell elongation was increased and final epidermal cell length was unchanged. These results showed that N deficiency reduced both cell division and cell elongation, which in turn resulted in decreased leaf length and prolonged time for leaf development. Nitrogen deficiency significantly increased both bulk and segmental cell wall peroxidase activity in the growth zone of both investigated genotypes, thus showing an interaction between leaf growth cessation and enzyme activity.     

Cuticular wax deposition in growing barley (<Emphasis Type="Italic">Hordeum vulgare</Emphasis>) leaves commences in relation to the point of emergence of epidermal cells from the sheaths of older leaves

In grasses, leaf cells divide and expand within the sheaths of older leaves, where the micro-environment differs from the open atmosphere. By the time epidermal cells are displaced into the atmosphere, they must have a functional cuticle to minimize uncontrolled water loss. In the present study, gas chromatography and scanning electron microscopy were used to follow cuticular wax deposition along the growing leaf three of barley (Hordeum vulgare L.). 1-Hexacosanol (C₂₆ alcohol) comprised more than 75% of extractable cuticular wax and was used as a marker for wax deposition. There was no detectable wax along the first 20 mm from the point of leaf insertion. Deposition started within the distal portion of the elongation zone (23–45 mm) and continued beyond the point of leaf emergence from the sheath of leaf two. The region where wax deposition commenced shifted towards more proximal (basal) positions when the point of leaf emergence was lowered by stripping back part of the sheath. When relative humidity in the shoot environment was elevated from 70% (standard growth conditions) to 92–96% for up to 4 days prior to analysis, wax deposition did not change significantly. The results show that cuticular waxes are deposited along the growing grass leaf independent of cell age or developmental stage. Instead, the reference point for wax deposition appears to be the point of emergence of cells into the atmosphere. The possibility of changes in relative humidity between enclosed and emerged leaf regions triggering wax deposition is discussed.     

Control of leaf cell elongation in barley. Generation rates of osmotic pressure and turgor, and growth-associated water potential gradients

In a previous study on the effects of N-supply on leaf cell elongation, the spatial distribution of relative cell elongation rates (RCER), epidermal cell turgor, osmotic pressure (OP) and water potential (Ψ) along the elongation zone of the third leaf of barley was determined (W. Fricke et al. 1997, Planta 202: 522–530). The results suggested that in plants receiving N at fixed relative addition rates (N-supply limitation of growth), cell elongation was rate-limited by the rate of solute provision, whereas in plants growing on complete nutrient solution containing excessive amounts of N (N-demand limitation), cell elongation was rate-limited by the rate of water supply or wall yielding. In the present paper, these suggestions were tested further. The generation rates of cell OP, turgor and Ψ along the elongation zone were calculated by applying the continuity equation of fluid dynamics to the previous data. To allow a more conclusive interpretation of results, anatomical data were collected and bulk solute concentrations determined. The rate of OP generation generally exceeded the rate of turgor generation. As a result, negative values of cell Ψ were created, particularly in demand-limited plants. These plants showed highest RCER along the elongation zone and a Ψ gradient of at least −0.15 MPa between water source (xylem) and expanding epidermal cells. The latter was similar to a theoretically predicted value (−0.18 MPa). Highest rates of OP generation were observed in demand-limited plants, with a maximum rate of 0.112 MPa · h⁻¹ at 16–20 mm from the leaf base. This was almost twice the rate in N-supply-limited plants and implied that the cells in the leaf elongation zone were capable of importing (or synthesising) every minute almost 1 mM of osmolytes. Potassium, Cl⁻ and NO₃ ⁻ were the main inorganic osmolytes (only determined for demand-limited plants). Their concentrations suggest that, unlike the situation in fully expanded epidermal cells, sugars are used to generate OP and turgor. Anatomical data revealed that the zone of lateral cell expansion extended distally beyond the zone of cell elongation. It is concluded that leaf cell expansion in barley relies on high rates of water and solute supply, rates that may not be sustainable during periods of sufficient N-supply (limitation by water supply: Ψ gradients) or limiting N-supply (limitation by solute provision: reduced OP-generation rates). To minimise the possibility of growth limitation by water and osmolyte provision, longitudinal and lateral cell expansion peak at different locations along the growth zone. Received: 15 October 1997 / Accepted: 12 March 1998     

Effects of Irradiance-Sulphur Interactions on Enzymes of Carbon,Nitrogen, and Sulphur Metabolism in Maize Plants

The effect of sulphur deprivation and irradiance (180 and 750 µmol m^–2 s^–1) on plant growth and enzyme activities of carbon, nitrogen, and sulphur metabolism were studied in maize (Zea mays L. Pioneer cv. Latina) plants over a 15-d-period of growth. Increase in irradiance resulted in an enhancement of several enzyme activities and generally accelerated the development of S deficiency. ATP sulphurylase (ATPs; EC 2.7.7.4) and o-acetylserine sulphydrylase (OASs; EC 4.2.99.8) showed a particular and different pattern as both enzymes exhibited maximum activity after 10 d from the beginning of deprivation period. Hence in maize leaves the enzymes of C, N, and S metabolism were differently regulated during the leaf development by irradiance and sulphur starvation.     

Age and growth of Pacific herring larvae based on length-frequency analysis and otolith ring number

Synopsis Age and growth in length and dry weight of cohorts of wild Pacific herring larvae, Clupea harengus pallasi, were measured using successive modes in the length-frequencies of the catches and the number of rings in the otoliths. Average linear rates of growth in length ranged from 0.36–0.41 mm d^–1, and average exponential rates of growth in dry weight ranged from 0.063–0.084 d^–1. Length-date and dry weight-age curves were best described using one-cycle and two-cycle Gompertz functions, respectively. Weight-length relationship were, therefore, curvilinear on double logarithmic plots and were best described by a non-linear allometric function. Average rates of otolith ring deposition were 0.90, 1.09 and 0.73–1.26d^–1. Rings were deposited daily from the day of complete yolk absorption in the first two cohorts, but interrupted ring deposition was observed over the first 27 d of the third cohort. Relatively low water temperatures, <9° C, coincided with the interruption and may have caused it.     

Effect of crop residues on root growth and phosphorus acquisition of pearl millet in an acid sandy soil in Niger

The effect of long-term (1983–1988) applications of crop residues (millet straw, 2–4 t ha^-1 yr^–1) and/or mineral fertilizer (30 kg N, 13 kg P and 25 kg K ha^-1 yr^-1) on uptake of phosphorus (P) and other nutrients, root growth and mycorrhizal colonization of pearl millet (Pennisetum glaucum L.) was examined for two seasons (1987 and 1988) on an acid sandy soil in Niger. Treatments of the long-term field experiment were: control (–CR–F), mineral fertilizer only (–CR+F), crop residues only (+CR–F), and crop residues plus mineral fertilizer (+CR+F).In both years, total P uptake was similar for +CR–F and –CR+F treatments (1.6–3.5 kg P ha^-1), although available soil P concentration (Bray I P) was considerably lower in +CR–F (3.2 mg P kg^-1 soil) than in –CR+F (7.4) soil. In the treatments with mineral fertilizers (–CR+F; +CR+F), crop residues increased available soil P concentrations (Bray I P) from 7.4 to 8.9 mg kg^-1 soil, while total P uptake increased from 3.6 to 10.6 kg P ha^-1. In 1987 (with 450 mm of rainfall), leaf P concentrations of 30-day-old millet plants were in the deficiency range, but highest in the +CR+F treatment. In 1988 (699 mm), leaf P concentrations were distinctly higher, and again highest in the +CR+F treatment. In the treatments without crop residues (–CR–F; –CR+F), potassium (K) concentrations in the leaves indicated K deficiency, while application of crop residues (+CR–F; +CR+F) substantially raised leaf K concentrations and total K uptake. Leaf concentrations of calcium (Ca) and magnesium (Mg) were hardly affected by the different treatments.In the topsoil (0–30 cm), root length density of millet plants was greater for +CR+F (6.5 cm cm^-3) than for +CR–F (4.5 cm cm^-3) and –CR+F (4.2 cm cm^-3) treatments. Below 30 cm soil depth, root length density of all treatments declined rapidly from about 0.6 cm cm^-3 (30–60 cm soil depth) to 0.2 cm cm^-3 (120–180 cm soil depth). During the period of high uptake rates of P (42–80 DAP), root colonization with vesicular-arbuscular mycorrhizal (VAM) fungi was low in 1987 (15–20%), but distinctly higher in 1988 (55–60%). Higher P uptake of +CR+F plants was related to a greater total root length in 0–30 cm and also to a higher P uptake rate per unit root length (P influx). Beneficial effects of crop residues on P uptake were primarily attributed to higher P mobility in the soil due to decreased concentrations of exchangeable Al, and enhancement of root growth. In contrast, the beneficial effect of crop residues on K uptake was caused by direct K supply with the millet straw.     

Growth Pattern and Movement of Epidermal Cells within Leaves of Asphodelus tenuifolius Cav.

The pattern of growth (velocity field) in the intercalary growthzones of monocotyledon leaves can be determined from patternsof cell number density (number per unit length of cell file)and leaf elongation rates using theory based on a cell numberconservation equation. The case where elongation rate is non-steadywhile the pattern of cell number density is steady is discussedand a method for extending calculations into the meristem usingobservations of numbers of mitotic cells is outlined. Applicationof these methods is illustrated using data for epidermal cellsin the first leaf of Asphodelus tenuifolius Cav. During earlyleaf development, leaf elongation rate increased exponentiallybut cell number density and mitotic number density were steady.Cells 0.1 mm from the base of the leaf when leaves were 3.2mm long took 8.3 d to move through the growth zone. In leavesthat were 4 d older, similar cells took 5.1 d to traverse thegrowth zone. Increases in the rates of leaf elongation and ofcell movement appeared to be associated mainly with increasesin total rates of cell production in the epidermal meristem. Asphodelus tenuifolius Cav., Asphodelus fistulosus L., velocity field, meristem, mitotic cell number density, extension-only zone     

Why do leaves and leaf cells of N-limited barley elongate at reduced rates?

The objective of the present study was to assess whether, in barley, nitrogen supply limits the rate of leaf elongation through a reduction in (relative) cell elongation rate and whether this is attributable to a reduced turgor, a reduced availability of osmolytes or, by implication, changed wall properties. Plants were grown on full-strength Hoagland solution (“Hoagland”-plants), or on N-deficient Hoagland solution while receiving N at a relative addition rate of 16 or 8% N · plant-N⁻¹ · d⁻¹ (“16%-” and “8%-plants”). Hoagland-plants were demand-limited, whereas 16%- and 8%-plants were supply-limited in N. Third leaves were analysed for leaf elongation rate and final epidermal cell length, and, within the basal growing region, for the spatial distribution of relative segmental elongation rates (RSER, pin-pricking method), epidermal cell turgor (cell-pressure probe), osmotic pressure (OP, picolitre osmometry) and water potential (Ψ). During the development of the third leaf, plants grew at relative growth rates (relative increase in fresh weight ) of 18.2, 15.6 and 8.1% · d⁻¹ (Hoagland-, 16%- and 8%-plants, respectively). Final leaf length and leaf elongation rate were highest in Hoagland plants (ca. 34.1 cm and 2.33–2.60 mm · h⁻¹, respectively), intermediate in 16%- plants (31.0 cm and 1.89–1.96 mm · h⁻¹) and lowest in 8%-plants (29.4 cm and 1.41–1.58 mm · h⁻¹). These differences were accompanied by only small differences in final cell length, but large differences in cell-flux rates (146, 187 and 201 cells · cell-file⁻¹ · d⁻¹ in 8%-, 16%- and Hoagland-plants, respectively). The length of the growth zone (32–38 mm) was not much affected by N-levels (and nutrient technique). A decrease in RSER in the growth zone distal to 10 mm produced the significant effect of N-levels on leaf elongation rate. In all treatments, cell turgor was almost constant throughout the growing region, as were cell OP and Ψ in 16%- and 8%-plants. In Hoagland-plants, however, cell OP increased by ca. 0.1 MPa within the zone of highest elongation rates and, as a consequence, cell Ψ decreased simultaneously by 0.1 MPa. Cell Ψ increased considerably where elongation ceased. Within the zone where differences in RSERs were highest between treatments (10–34 mm from base) average turgor was lowest, OP highest and Ψ most negative in Hoagland- compared to 8%- and 16%-plants (P < 0.001), but not significantly different between 8%- and 16%-plants. Received: 9 January 1997 / Accepted: 6 March 1997     

Axial rod growth and age estimation of the sea pen,Halipteris willemoesi Kölliker

Halipteris willemoesi is a large octocoral commonly found in the Bering Sea. It is a member of a ubiquitous group of benthic cnidarians called sea pens (Octocorallia: Pennatulacea). Sea pens have a skeletal structure, the axial rod, that in cross section exhibits growth rings. Pairs of adjacent rings, or ring couplets, were assumed to be annuli and were used to estimate the age and growth of H. willemoesi. Twelve axial rods, extracted from H. willemoesi collected in the Bering Sea, were selected to represent small (25–29 cm total length), medium (97–130 cm TL) and large (152–167 cm TL) colonies. Each rod resembled a tapered dowel; the thickest part (0.90–6.75 mm in diameter) was at about 5–10% of total length from the base tip, the distal part was more gradually tapered than was the base. The number of ring couplets increased with rod size indicating their utility in estimating age and growth. Estimated age among rods was based on couplet counts at the thickest part of each rod; the average estimated age (±SE) was 7.1 ±0.7, 19.3 ±0.5, and 44.3 ±2.0 yr for small, medium and large-size rods, respectively. Based on these estimated ages, average growth rate in total length was 3.9 ± 0.2, 6.1 ± 0.3, and 3.6 ± 0.1 cm yr^–1 for small, medium, and large-size colonies. The average annual increase in maximum rod diameter among all colonies was 0.145 ± 0.003 SE mm yr^–1; therefore, age prediction from maximum rod diameter was calculated (estimated age (yr) = 7.0 * (maximum rod diameter, mm) –0.2; R ² = 0.99). At maximum diameter, the average couplet width was relatively constant among the three colony sizes (0.072 ± 0.05 mm). X-ray diffraction and electron microprobe analyses revealed that the inorganic portion of the rod is composed of a high-magnesium calcite. Radiometric validation of these age and growth rate estimates was attempted, but high amounts of exogenous ²¹⁰Pb precluded using the disequilibria of ²¹⁰Pb:²²⁶Ra. Instead, ²¹⁰Pb activities were measured in a series of cores extracted along the axial rod. These activities ranged from 0.691 ± 0.036 (SE) to 2.76 ± 0.13 dpm g^–1, but there was no pattern of decay along the length of the rod; therefore, the growth rates and corresponding ages could not be validated. Based on estimated age from ring couplet counts, growth in total rod length is slow at first, fastest at medium size, and slows toward maximum size, with an estimated longevity approaching 50 yr.     

Rapid and tissue-specific accumulation of solutes in the growth zone of barley leaves in response to salinity

The aim of the present study was to test whether rapid accumulation of solutes in response to salinity in leaf tissues of barley (Hordeum vulgare L.) contributes to recovery and maintenance of residual elongation growth. Addition of 100 mM NaCl to the root medium caused an immediate reduction close to zero in elongation velocity of the growing leaf 3. After 20–30 min, elongation velocity recovered suddenly, to 40–50% of the pre-stress level. Bulk osmolality increased first, after 60 min, significantly in the proximal half of the elongation zone. Over the following 3 days, osmolality increases became significant in the distal half of the elongation zone, the adjacent, enclosed non-elongation zone and finally in the emerged portion of the blade. The developmental gradient and time course in osmolality increase along the growing leaf was reflected in the pattern of solute (Cl, Na and K) accumulation in bulk tissue and epidermal cells. The partitioning of newly accumulated solutes between epidermis and bulk tissue changed with time. Even though solute accumulation does not contribute to the sudden and partial growth recovery 20–30 min after exposure to salt, it does facilitate residual growth from 1 h onwards. This is due to a high sink strength for solutes of the proximal part of the growth zone and its ability to accumulate solutes rapidly and at high rates.Abbreviations EDX analysis Energy-dispersive X-ray analysis - LEV Leaf elongation velocity - LVDT Linear variable differential transformer - REGR Relative elemental growth rate     

Carbon and nitrogen deposition in expanding tissue elements of perennial ryegrass (Lolium perenne L.) leaves during non-steady growth after defoliation

The effect of defoliation on the deposition of carbon (C) and nitrogen (N) and the contribution of reserves and current assimilates to the use of C and N in expanding leaf tissue of severely defoliated perennial ryegrass (Lolium perenne L.) was assessed with a new material element approach. This included ¹³C/¹²C-and ¹⁵N/¹⁴N-steady-state labelling of all post-defoliation assimilated C and N, analysis of tissue expansion and displacement in the growth zone, and investigation of the spatial and temporal changes in substrate and label incorporation in the expanding elements prior to and after defoliation. The relationship between elemental expansion and C deposition was not altered by defoliation, but total C deposition in the growth zone was decreased due to decreased expansion of tissue at advanced developmental stages and a shortening of the growth zone. The N deposition per unit expansion was increased following defoliation, suggesting that N supply did not limit expansion. Transition from reserve- to current assimilation-derived growth was rapid (<1 d for carbohydrates and approximately 2 d for N), more rapid than suggested by label incorporation in growth zone biomass. The N deposition was highest near the leaf base, where cell division rates are greatest, whereas carbohydrate deposition was highest near the location of most active cell expansion. The contribution of reserve-derived relative to current assimilation-derived carbohydrates (or N) to deposition was very similar for elements at different stages of expansion     

The involvement of the root apex and cytokinins in the control of lateral root emergence in lettuce seedlings

Removal of the apical 3 mm of the primary root of hydroponically-grown lettuce seedlings 3 or 5 days after sowing, prevented further elongation of the root and increased both the number and total length of lateral roots. The length of the lateral zone, i.e. the distance from the base of the parent root to the lateral nearest the tip, except on one occasion, remained the same as the control in both 3 and 5 day treatments, until the length of the decapitated root (which had ceased elongating) became limiting.Zeatin applied via the roots, at a concentration range from 3 × 10^–10 M to 10^–8 M reduced tap root extension growth at all concentrations. Lateral root emergence was enhanced by low zeatin concentrations and retarded by higher ones. In general, the lateral zone length was the same in cytokinin-treated plants as in untreated controls.     

Dynamics of leaf minerals,bdleaf area,and biomass from hardwoods intensively grown on a peat bog

Summary Water willow and grey alder were grown on a raised sphagnum bog in central Sweden. The stands were intensively treated by daily irrigation and fertilization during the growing period in order to improve site fertility. After a 2-year establishment period high production rates were achieved in willow stands, 0.8 kg stem dry weight m^–2 year^–1 on current plus one (C+1) year old shoots. In these stands the canopy was closed with a leaf area index (LAI) that peaked at approximately 7. The canopy in the alder stand did not close during the initial 3 years of growth and the measured production rate was relatively low, at approximately 0.4 kg dry weight m^–2 year ^–1 in the last year. The leaf nitrogen content was 3%–4% of dry weight during the summer and the other studied mineral elements were in almost optimal proportion to nitrogen. This was considered to be an effect of the intensive fertilization regime. Above-ground production close to maximum yield was attained in the prevailing conditions of soil, climate and biomass partitioning.     

Rapid nutrient cycling in leaf litter from invasive plants in Hawai’i

Physiological traits that contribute to the establishment and spread of invasive plant species could also have impacts on ecosystem processes. The traits prevalent in many invasive plants, such as high specific leaf areas, rapid growth rates, and elevated leaf nutrient concentrations, improve litter quality and should increase rates of decomposition and nutrient cycling. To test for these ecosystem impacts, we measured initial leaf litter properties, decomposition rates, and nutrient dynamics in 11 understory plants from the Hawaiian islands in control and nitrogen + phosphorus fertilized plots. These included five common native species, four of which were ferns, and six aggressive invasive species, including five angiosperms and one fern. We found a 50-fold variation in leaf litter decay rates, with natives decaying at rates of 0.2–2.3 year^–1 and invaders at 1.4–9.3 year^–1. This difference was driven by very low decomposition rates in native fern litter. Fertilization significantly increased the decay rates of leaf litter from two native and two invasive species. Most invasive litter types lost nitrogen and phosphorus more rapidly and in larger quantities than comparable native litter types. All litter types except three native ferns lost nitrogen after 100 days of decomposition, and all litter types except the most recalcitrant native ferns lost >50% of initial phosphorus by the end of the experiment (204–735 days). If invasive understory plants displace native species, nutrient cycling rates could increase dramatically due to rapid decomposition and nutrient release from invasive litter. Such changes are likely to cause a positive feedback to invasion in Hawaii because many invasive plants thrive on nutrient-rich soils.     

Nitrogen fixation and metabolism by groundwater-dependent perennial plants in a hyperarid desert

The Central Asian Taklamakan desert is characterized by a hyperarid climate with less than 50 mm annual precipitation but a permanent shallow groundwater table. The perched groundwater (2–16 m) could present a reliable and constant source of nitrogen throughout the growing season and help overcome temporal nitrogen limitations that are common in arid environments. We investigated the importance of groundwater and nitrogen fixation in the nitrogen metabolism of desert plants by assessing the possible forms and availability of soil N and atmospheric N and the seasonal variation in concentration as well as isotopic composition of plant N. Water availability was experimentally modified in the desert foreland through simulated flooding to estimate the contribution of surface water and temporally increased soil moisture for nutrient uptake and plant–water relations. The natural vegetation of the Taklamakan desert is dominated by plants with high foliar nitrogen concentrations (2–3% DM) and leaf nitrate reductase activity (NRA) (0.2–1 mol NO₂^– g^–1 FW h^–1). There is little evidence that nitrogen is a limiting resource as all perennial plants exhibited fast rates of growth. The extremely dry soil conditions preclude all but minor contributions of soil N to total plant N so that groundwater is suggested as the dominant source of N with concentrations of 100 M NO₃^–. Flood irrigation had little beneficial effect on nitrogen metabolism and growth, further confirming the dependence on groundwater. Nitrogen fixation was determined by the ¹⁵N natural abundance method and was a significant component of the N-requirement of the legume Alhagi, the average contribution of biologically fixed nitrogen in Alhagi was 54.8%. But nitrogen fixing plants had little ecological advantage owing to the more or less constant supply of N available from groundwater. From our data we conclude that the perennial species investigated have adapted to the environmental conditions through development of root systems that access groundwater to satisfy demands for both water and nutrients. This is an ecologically favourable strategy since only groundwater is a predictable and stable resource.     

Warming chambers stimulate early season growth of an arctic sedge: results of a minirhizotron field study

We examined the effects of passive open-top warming chambers on Eriophorum vaginatum production near Toolik Lake, Alaska, USA. During the 2002 growing season, chamber warming was consistent with the magnitude and seasonality observed in recent decades throughout northwestern North America. Leaf-growth rates were higher in late May and early June; maximum growth rates in each leaf cohort occurred earlier and peak biomass was observed 20 days earlier within the chambers. Consequently, plants within the chambers maintained more live leaf biomass during the period of highest photosynthetically active radiation. Annual leaf production within the chambers (21±2 mg tiller) was not significantly different than under ambient conditions (17±2 mg tiller) (P=0.2256) despite higher early-season growth rates. Root growth began earlier; growth rates were higher in late May and early June, and maximum growth rates occurred earlier within the chambers. Therefore, plants within the chambers maintained greater root biomass during what earlier studies have identified as a period of relatively high nutrient availability. Annual root production within the chambers (191±42 g m^–2) was not significantly different than under ambient conditions (119±48 g m^–2) (P=0.1979), although there was a trend toward higher production within the chambers. The tendency toward higher root production within the chambers is consistent with previous laboratory experiments and with the predictions of biomass allocation theory.