The energetic importance of terrestrial arthropod inputs to three warm-water streams

1. Inputs of terrestrial arthropods (number and mass m^–2 d^–1) from riparian corridors to three streams representing different orders were highly variable among seasons and sites, with significantly greater ( P < 0.05) inputs at the headwater stream during summer months, compared with other sites and seasons.
2. No significant differences in estimates of stream retention of terrestrial arthropods (number and mass m^–2 d^–1) were observed among sites; however, retention of terrestrial arthropods at all sites was significantly greater during summer months, compared with other periods.
3. The gravimetric proportion of terrestrial arthropods present in the stomachs of redbreast sunfish ( Lepomis auritus ) and bluegill ( L. macrochirus ) was equivalent among sites. However, estimates of the dietary importance of terrestrial arthropods at all study sites were significantly greater in the summer, compared with other seasons.
4. Estimates of the potential annual energetic contribution (kJ m^–2 d^–1) of terrestrial arthropod inputs to the stream system were comparable with published rates of total annual production of aquatic macroinvertebrates in other Virginia headwater streams.
5. Results of this study supported the hypothesis that terrestrial arthropods represented an important energetic subsidy to stream fish during periods of low aquatic macroinvertebrate availability, and suggest that this component of allochthonous input is a potentially significant, but poorly understood energetic linkage between riparian landscapes and stream ecosystems.     

The energetic importance of terrestrial arthropod inputs to three warm-water streams

1. Inputs of terrestrial arthropods (number and mass m^–2 d^–1) from riparian corridors to three streams representing different orders were highly variable among seasons and sites, with significantly greater ( P < 0.05) inputs at the headwater stream during summer months, compared with other sites and seasons.
2. No significant differences in estimates of stream retention of terrestrial arthropods (number and mass m^–2 d^–1) were observed among sites; however, retention of terrestrial arthropods at all sites was significantly greater during summer months, compared with other periods.
3. The gravimetric proportion of terrestrial arthropods present in the stomachs of redbreast sunfish ( Lepomis auritus ) and bluegill ( L. macrochirus ) was equivalent among sites. However, estimates of the dietary importance of terrestrial arthropods at all study sites were significantly greater in the summer, compared with other seasons.
4. Estimates of the potential annual energetic contribution (kJ m^–2 d^–1) of terrestrial arthropod inputs to the stream system were comparable with published rates of total annual production of aquatic macroinvertebrates in other Virginia headwater streams.
5. Results of this study supported the hypothesis that terrestrial arthropods represented an important energetic subsidy to stream fish during periods of low aquatic macroinvertebrate availability, and suggest that this component of allochthonous input is a potentially significant, but poorly understood energetic linkage between riparian landscapes and stream ecosystems.     

Non-photosynthetic mechanisms of growth reduction in pea (Pisum sativum L.) exposed to UV-B radiation

Pisum sativum cv. Guido grown under controlled environment conditions was exposed to either low or high UV-B radiation (2·2 or 9·9 kJ m^–2 d^–1 plant-weighted UV-B, respectively). Low or high UV-B was maintained throughout growth (LL and HH treatments, respectively) or plants were transferred between treatments when 22 d old (giving LH and HL treatments). High UV-B significantly reduced plant dry weight and significantly altered plant morphology. The growth and morphology of plants transferred from low to high UV-B were little affected, when compared with those of LL plants. By contrast, plants moved from high to low UV-B showed marked increases in growth when compared with HH plants. This contrast between HL and LH appeared to be related to the effect of UV-B on plant development. Exposure to high UV-B throughout development consistently reduced leaf areas. In fully expanded leaves there was no significant UV-B effect on cell area and reduced leaf area could be attributed to reduced cell number, suggesting effects on leaf primordia. Further reductions in the leaf area of younger leaves were the result of the slower development rate of plants grown at high UV-B, which also resulted in significant reductions in leaf number.     

Pea seedling growth and development regulated by cotyledons and modified by irradiance

Growth and development of hydroponically grown pea seedlings ( Pisum sativum L. cv. Alaska) were measured using stem and root length as well as number of leaves and lateral roots. The growth was dependent on the presence of cotyledons and was modulated by the irradiance. All plants were grown in a full nutrient solution. If grown at low irradiance (73 μmol m^-2s^-1) they depended more and for a longer time on the cotyledons than plants grown at high irradiance (220 μmol m^-2s^-1). Low irradiance caused stem elongation but decreased root length and number of lateral roots as compared to plants grown at high irradiance. The dark respiration of the leaves was measured as oxygen uptake. In plants grown at the low irradiance, excision of the cotyledons caused the rate of oxygen uptake to increase by a factor of three, and the increase was sensitive to cyanide. Decotyledonized plants showed a high respiration rate and a diminished leaf growth for their entire life cycle. CO₂ fixation also increased in decotyledonized pea seedlings grown at either irradiance. The mobilization of food reserves from the seeds was positively correlated to seed dry weight, but only if the plants were grown at 73 μmol m^-2s^-1. Increasing dry weight of the seed enhanced top growth, whereas root growth was depressed, so that top and root responds differently with regard to that part of growth which depends on mobilization of reserves from the seed.     

The effects of light on induction, time courses, and kinetic patterns of net nitrate uptake in barley

Barley seedlings ( Hordeum vulgare L.) were grown hydroponically with (induced) or without (uninduced) nitrate in a light/dark cycle with high photon flux density to determine the effects of light on time courses, induction and kinetics of net nitrate uptake. Nitrate uptake was induced by external nitrate in both light and dark and was prevented by 1 mol m^–3 p-fluorophenylalanine. In high light, nitrate uptake was about 2-fold higher than in low light. During time course experiments the uptake rates oscillated due to daily light–dark changes. Rates of nitrate uptake also increased at about 2200 h during continuous darkness. This increase coincided approximately with the time at which the dark period started during the previous culture of the plants, indicating that it was due to a mechanism associated with an endogenous diurnal rhythm. When calculating the kinetics of nitrate uptake, a model with two saturable systems, including a high-affinity system (HATS) and a low-affinity system (LATS), gave the best fit to data in all treatments. The apparent affinity of the HATS ranged from 7·7 to 12·2 mmol m^–3 in induced plants in all light conditions. The effect of light on the HATS was mainly an increase of apparent V _max in the step from low to high light. In uninduced plants the HATS operated at a very low activity which was strongly enhanced during induction. Interpretation of the calculated kinetics of the LATS was much more difficult on the basis of net uptake data. The apparent affinity of the LATS increased from 24·3 mol m^–3 in low light up to 0·17 mol m^–3 after acceleration in high light. These extreme changes in apparent affinity of the LATS could not be explained satisfactorily, and the nature of this system is also discussed with respect to the method used.     

The consequences of freezing temperatures followed by high irradiance on in vivo chlorophyll fluorescence and growth in Picea abies

Picea abies (L.) Karst. plants, propagated by cuttings, were subjected to one night of freezing temperatures (-5°C), high irradiance (1 200 or 1 800 μmol m⁻² s⁻¹), or freezing temperatures followed by high irradiance. The treatments were applied at bud burst, at time of shoot elongation, and when the shoots had ceased to elongate. The maximum quantum yield of photosynthesis, F_v/F_m, dry weight of branches and needles, and length and survival of shoots were measured. F_v/F_m and growth decreased after a night of freezing temperatures followed by high irradiance, at the time of bud burst and shoot elongation. High irradiance alone influenced F_v/F_m, but not growth. Freezing temperatures affected F_v/F_m, and growth at the time of shoot elongation. F₀ increased after a night of freezing temperatures and decreased with age of the current-year needles. It was concluded that the use of short-term measurements of chlorophyll fluorescence induction to predict changes in growth after a night of frost and subsequent high light was not a reliable method.     

Changes in pigment levels, Rubisco and respiratory enzyme activity of Ficus benjamina during acclimation to low irradiance

The objective of the present study was to determine the influence of reduced irradiance on the activities of ribulose bisphosphate carboxylase-oxygenase (Rubisco) and respiratory enzymes. Rooted cuttings of the tropical epiphyte. Ficus benjamina L., were grown in a shaded environment that excluded approximately 50% of the natural photosynthetically active irradiance (890 μmol m⁻² s⁻¹) for 4 months. Established plants were transferred and grown for 10 months under a range of irradiance levels with daily average maxima varying from a full-sun environment to 20% full sun (100%−1735; 50%−890; 40%−695; and 20%−303 μmol m⁻²s⁻¹). Chlorophyll, carotenoid and soluble protein content increased in Ficus leaves as irradiance level decreased, while Rubisco increased on a fresh weight basis but decreased on a protein basis. Glycolytic enzymes, enolase and pyruvate kinase, showed higher activities in full-sun plants on a protein and fresh weight basis. However, the activity of two mitochondrial enzymes, aconitase and malate dehydrogenase, was not different under the various irradiance levels. When transferred to a very low irradiance environment (18 μmol m⁻² s⁻¹), mature leaves exhibited increased chlorophyll and carotenoid levels regardless of previous irradiance treatment. Exposure to very low irradiance resulted in a large increase in enolase and pyruvate kinase activities. Only plants grown under full sun conditions showed a decline in Rubisco activity following growth at very low irradiance. Together, these studies demonstrate the ability of mature leaves of Ficus to biochemically adjust photosynthetic and respiratory components over a wide range of irradiance.     

Acquisition and allocation of carbon and nitrogen by Danthonia richardsonii in response to restricted nitrogen supply and CO2 enrichment

Dry weight (DW) and nitrogen (N) accumulation and allocation were measured in isolated plants of Danthonia richardsonii (Wallaby Grass) for 37 d following seed imbibition. Plants were grown at ≈ 365 or 735 μ L L^–1 CO₂ with N supply of 0·05, 0·2 or 0·5 mg N plant^–1 d^–1. Elevated CO₂ increased DW accumulation by 28% (low-N) to 103% (high-N), following an initial stimulation of relative growth rate. Net assimilation rate and leaf nitrogen productivity were increased by elevated CO₂, while N concentration was reduced. N uptake per unit root surface area was unaffected by CO₂ enrichment. The ratio of leaf area to root surface area was decreased by CO₂ enrichment. Allometric analysis revealed a decrease in the shoot-N to root-N ratio at elevated CO₂, while the shoot-DW to root-DW ratio was unchanged. Allometric analysis showed leaf area was reduced, while root surface area was unchanged by elevated CO₂, indicating a down-regulation of total plant capacity for carbon gain rather than a stimulation of mineral nutrient acquisition capacity. Overall, growth in elevated CO₂ resulted in changes in plant morphology and nitrogen use, other than those associated simply with changing plant size and non-structural carbohydrate content.     

Gas exchange and carbon isotope composition of Ananas comosus in response to elevated CO2 and temperature

Ananas comosus L. (Merr.) (pineapple) was grown at three day/night temperatures and 350 (ambient) and 700 (elevated) μ mol mol^–1 CO₂ to examine the interactive effects of these factors on leaf gas exchange and stable carbon isotope discrimination ( Δ ,‰). All data were collected on the youngest mature leaf for 24 h every 6 weeks. CO₂ uptake (mmol m^–2 d^–1) at ambient and elevated CO₂, respectively, were 306 and 352 at 30/20 °C, 175 and 346 at 30/25 °C and 187 and 343 at 35/25 °C. CO₂ enrichment enhanced CO₂ uptake substantially in the day in all environments. Uptake at night at elevated CO₂, relative to that at ambient CO₂, was unchanged at 30/20 °C, but was 80% higher at 30/25 °C and 44% higher at 35/25 °C suggesting that phosphoenolpyruvate carboxylase was not CO₂-saturated at ambient CO₂ levels and a 25 °C night temperature. Photosynthetic water use efficiency (WUE) was higher at elevated than at ambient CO₂. Leaf Δ -values were higher at elevated than at ambient CO₂ due to relatively higher assimilation in the light. Leaf Δ was significantly and linearly related to the fraction of total CO₂ assimilated at night. The data suggest that a simultaneous increase in CO₂ level and temperature associated with global warming would enhance carbon assimilation, increase WUE, and reduce the temperature dependence of CO₂ uptake by A. comosus .     

The stomatal response to CO2 is linked to changes in guard cell zeaxanthin*

The mechanisms mediating CO₂ sensing and light–CO₂ interactions in guard cells are unknown. In growth chamber-grown Vicia faba leaves kept under constant light (500 μ mol m^–2 s^–1) and temperature, guard cell zeaxanthin content tracked ambient [CO₂] and stomatal apertures. Increases in [CO₂] from 400 to 1200 cm³ m^–3 decreased zeaxanthin content from 180 to 80 mmol mol^–1 Chl and decreased stomatal apertures by 7·0 μ m. Changes in zeaxanthin and aperture were reversed when [CO₂] was lowered. Guard cell zeaxanthin content was linearly correlated with stomatal apertures. In the dark, the CO₂-induced changes in stomatal aperture were much smaller, and guard cell zeaxanthin content did not change with chamber [CO₂]. Guard cell zeaxanthin also tracked [CO₂] and stomatal aperture in illuminated stomata from epidermal peels. Dithiothreitol (DTT), an inhibitor of zeaxanthin formation, eliminated CO₂-induced zeaxanthin changes in guard cells from illuminated epidermal peels and reduced the stomatal CO₂ response to the level observed in the dark. These data suggest that CO₂-dependent changes in the zeaxanthin content of guard cells could modulate CO₂-dependent changes of stomatal apertures in the light while a zeaxanthin-independent CO₂ sensing mechanism would modulate the CO₂ response in the dark.     

In situ responses to elevated CO2 in tropical forest understorey plants

1. Plants growing in deep shade and high temperature, such as in the understorey of humid tropical forests, have been predicted to be particularly sensitive to rising atmospheric CO₂. We tested this hypothesis in five species whose microhabitat quantum flux density (QFD) was documented as a covariable. After 7 (tree seedlings of Tachigalia versicolor and Beilschmiedia pendula ) and 18 months (shrubs Piper cordulatum and Psychotria limonensis, and grass Pharus latifolius ) of elevated CO₂ treatment ( c. 700 μl litre^–1) under mean QFD of less than 11 μmol m^–2 s^–1, all species produced more biomass (25–76%) under elevated CO₂.
2. Total plant biomass tended to increase with microhabitat QFD (daytime means varying from 5 to 11μmol m^–2 s^–1) but the relative stimulation by elevated CO₂ was higher at low QFD except in Pharus .
3. Non-structural carbohydrate concentrations in leaves increased significantly in Pharus (+ 27%) and Tachigalia (+ 40%).
4. The data support the hypothesis that tropical plants growing near the photosynthetic light compensation point are responsive to elevated CO₂. An improved plant carbon balance in deep shade is likely to influence understorey plant recruitment and competition as atmospheric CO₂ continues to rise.     

Diurnal changes in phosphoenolpyruvate carboxylase and pyruvate, orthophosphate dikinase properties in the natural environment: interplay of light and temperature in a C4 thermophile

Activities of phosphoenolpyruvate (PEP) carboxylase (EC 4.1.1.31) were measured in leaf extracts of field grown Amaranthus paniculatus L. (C₄) during a natural diurnal irradiance and temperature pattern. Enzyme assays were run at both fixed (30°C) and the corresponding leaf temperature at the time of harvest. Light activation of PEP carboxylase (PEPCase) at fixed assay temperatures was expressed as a decrease in S_0–5 (PEP) after a threshold (> 330 μmol m^–2 s^–1) photon fluence rate was surpassed at noon. Earlier in the morning, increase in apparent enzyme affinity for PEP was observed when the assay was run at leaf temperature, indicating a physiologically meaningfull effect of temperature on S^0.5 (PEP). The 3.3-fold increase in PEPCase activity at low PEP and fixed assay temperature between the minimal and maximal irradiance and temperature hours of the day, became 12.8-, 11.5- and 7.4-fold when assays were run at the corresponding leaf temperature during three diurnal cycles with respective temperature differences (max minus min) of 9.0, 8.3 and 7.4°C. The extent of malate inhibition was the same for both day and night forms of PEPCase assayed at 35°C, but increased considerably with night enzyme at 25°C. The results indicate that light increases the apparent affinity of PEPCase for PEP and that at lower temperatures malate becomes more inhibitory. Pyruvate orthophosphate dikinase activity started to increase immediately after sunrise and the 10-fold increase at fixed temperature became 14.8-, 14.2- and 13.1-fold when assays were run at the above leaf temperatures. This indicates that the light effect predominates with pyruvate, orthophosphate dikinase, while with phosphoenolpyravate carboxylase, light and temperature co-operate to increase the day enzyme activities.     

Long-term effects of successive Ca(OH)2 and CaCO3 treatments on the water quality of two eutrophic hardwater lakes

1. Whole-lake experiments were conducted in two hardwater lakes (Halfmoon and Figure Eight) in Alberta, Canada, to investigate the effectiveness of repeated lime (slaked lime: Ca(OH)₂ and/or calcite: CaCO₃) treatments (5–78 mg L^–1) for up to 7 years.
2. Randomized intervention analysis of intersystem differences between the experimental and three reference lakes demonstrated a decline in euphotic total phosphorus and chlorophyll a concentrations in the experimental lakes after repeated lime treatments.
3. After the second lime application to Halfmoon Lake, mean winter total phosphorus release rates (TPRR) decreased to < 1 mg m^–2 day^–1 compared with 3.6 mg m^–2 day^–1 during the winter after initial treatment. In the final year of lime application, mean summer TPRR decreased to 4.5 mg m^–2 day^–1 compared with 7.6 mg m^–2 day^–1 in the pre-treatment year.
4. Mean macrophyte biomass declined and species composition was altered at 1 and 2 m depths in Figure Eight Lake during lime application. Over the first 6 years of treatment, macrophyte biomass at 2 m declined by 95% compared with concentrations recorded during the initial treatment year. In the last year of the study, macrophyte biomass at 2 m reached initial treatment concentrations, which coincided with the greatest water transparency. Over the treatment period, macrophyte species shifted from floating to rooted plants.
5. Multiple lime applications can improve water quality in eutrophic hardwater lakes for periods of up to 7 years.     

Particulate organic matter inputs to a glacial stream ecosystem in the Swiss Alps

1. Traps for litterfall and for lateral transport of organic matter were sampled over a 1-year period along longitudinal and lateral transects in a glacial stream system (Val Roseg, Swiss Alps), which is characterized by single-thread reaches and a large subalpine floodplain.
2. Allochthonous inputs to the glacier stream were low close to the glacier terminus but increased as woody riparian vegetation and forests develop. Annual inputs varied from 0.4 g ash free dry matter (AFDM) m^–2 year^–1 (direct input) and 0.7 g AFDM m^–2 year^–1 (lateral input) in the proglacial area to 23.0 g AFDM m^–2 year^–1 (direct input) and 10.7 g AFDM m^–2 year^–1 (lateral input) in the lowest reach with adjacent subalpine forests.
3. Direct inputs of organic matter decreased exponentially from forests at the floodplain edge to the floodplain centre, while lateral inputs of organic matter correlated linearly with distance to trees. Direct litterfall dominated litter input close to the forest, while lateral transport was the major pathway for channels more than 20 m away from the forest.
4. A conceptual framework is developed illustrating the influence of terrestrial vegetation and fluvial morphology on organic matter input along the continuum of glacial streams.     

Elevated atmospheric CO2 does not affect per se the preference for symbiotic nitrogen as opposed to mineral nitrogen of Trifolium repens L.

The objective of this investigation was to examine the effect of an elevated atmospheric CO₂ partial pressure ( p CO₂) on the N-sink strength and performance of symbiotic N₂ fixation in Trifolium repens L. cv. Milkanova. After initial growth under ambient p CO₂ in a nitrogen-free nutrient solution, T. repens in the exponential growth stage was exposed to ambient and elevated p CO₂ (35 and 60 Pa) and two levels of mineral N (N-free and 7·5 mol m^–3 N) for 36 d in single pots filled with silica sand in growth chambers. Elevated p CO₂ evoked a significant increase in biomass production from day 12 after the start of CO₂ enrichment. For plants supplied with 7·5 mol m^–3 N, the relative contribution of symbiotically fixed N (%N_sym) as opposed to N assimilated from mineral sources (¹⁵N-isotope-dilution method), dropped to 40%. However, in the presence of this high level of mineral N, %N_sym was unaffected by atmospheric p CO₂ over the entire experimental period. In plants fully dependent on N₂ fixation, the increase in N yield reflects a stimulation of symbiotic N₂ fixation that was the result of the formation of more nodules rather than of higher specific N₂ fixation. These results are discussed with regard to physiological processes governing symbiotic N₂ fixation and to the response of symbiotic N₂ fixation to elevated p CO₂ in field-grown T. repens .     

Overexpression of light-dependent PORA or PORB in plants depleted of endogenous POR by far-red light enhances seedling survival in white light and protects against photooxidative damage

The structurally related light-dependent protochlorophyllide (Pchlide) oxidoreductases PORA and PORB mediate the only light-requiring step in chlorophyll (Chl) biosynthesis in higher plants. Correlative evidence suggests that some in vivo functions of PORA and PORB may be unique, including a postulated photoprotective role for PORA. For example, wild-type Arabidopsis thaliana seedlings grown in non-photooxidative far-red light (cFR) resemble those grown in white light (WL), but they are yellow and do not green normally thereafter in WL. This defect is accompanied by the absence of detectable PORA and reduced levels of PORB expression. Here, direct evidence is provided that the presence of POR, either as PORA or PORB, can confer photoprotection in plants. In contrast to the wild-type, the plastids of transgenic PORA- or PORB-overexpressing Arabidopsis seedlings grown in cFR possess extensive prolamellar bodies. Upon a subsequent shift to WL, POR-overexpressing seedlings develop thylakoid membranes, accumulate large amounts of Chl and are viable at fluence rates lethal to the wild-type. Intriguingly, the plastid membrane architectures of greening transgenic seedlings seem to depend on whether PORA or PORB has been overproduced. POR-overexpressing seedlings shifted from cFR to WL of fluence rates from 20 to 500 μE m^–2 sec^–1 accumulate substantially higher amounts of Chl than does the wild-type. Furthermore, the WL fluence rate that permits maximal Chl accumulation increases from 8 μE m^–2 sec^–1 in the wild-type to 125 μE m^–2 sec^–1 in transgenic seedlings. POR overexpression during growth in cFR also correlates with a fourfold decrease in the steady-state content of Pchlide, a potentially lethal photosensitizer.     

Photosynthetic acclimation to elevated atmospheric carbon dioxide and UV irradiation in Pinus banksiana

Pinus banksiana seedlings were grown for 9 months in enclosures in greenhouses at CO₂ concentrations of 350 or 750 μmol mol⁻¹ with either low (0.005 to 0. 3 W m⁻²) or high (0.25 to 0. 90 W m⁻²) ultraviolet-B (UV-B) irradiances. Total seedling dry weight decreased with high UV treatment but was unaffected by CO₂ enrichment. High UV treatment also shifted biomass partitioning in favor of leaf production. Both CO₂ and UV treatments decreased the dark respiration rate and light compensation point. High UV light inhibited photosynthesis at 350 but not at 750 μmol mol⁻¹ CO₂ due to a UV induced increase in ribulose-1, 5-bisphosphate carboxylase/oxygenase efficiency and ribulose-1, 5-bisphosphate regeneration. Stomatal density was increased by high UV irradiance but was unchanged by CO₂ enrichment.     

Low irradiance modifies the effect of water stress on survival and growth-related parameters during the early developmental stages of buckwheat (Fagopyrum esculentum)

In order to characterize physiological modifications encountered by buckwheat plants exposed to both drought and low-light stresses, seedlings (cv. La Harpe) were exposed under controlled environmental conditions, to a progressive decline in soil volumetric water content under two light regimes: low irradiance (80 µmol m⁻² s⁻¹) or moderate irradiance (160 µmol m⁻² s⁻¹). Phenological evolution of the whole plant until the macroscopic appearance of the reproductive structure and physiological properties of leaves in relation to their position on the main axis were quantified. Water stress reduced net assimilation rate (NAR) before specific leaf area (SLA) and induced a decrease in stomatal conductance (g_l) and carbon isotope discrimination (Δ). Water consumption by stressed plants was similar under both light treatments. Water-stressed plants under moderate irradiance exhibited higher growth, NAR, osmotic adjustment, and lower SLA than plants maintained under low irradiance. However, the former died after 27 days of treatment while the latter still remained alive until the experiment was discontinued (40 days). We concluded that the physiological strategy adopted by the water-stressed plants maintained under moderate irradiance did not afford a long-term advantage in terms of survival. The effects of a combination of low-light and water stress on chlorophyll concentration and carbon isotope discrimination (Δ) are discussed in relation to growth parameters.     

Effect of phosphorus supply on the formation and function of proteoid roots of white lupin (Lupinus albus L.)

We investigated (1) the effect of constant and altered inorganic phosphate (P_i) supply (1–100 mmol m^–3) on proteoid root production by white lupin ( Lupinus albus L.); and (2) the variation in citrate efflux, enzyme activity and phosphate uptake along the proteoid root axis in solution culture. Proteoid root formation was greatest at P_i solution concentrations of 1–10 mmol m^–3 and was suppressed at 25 mmol m^–3 P_i and higher. Except at 1 mmol m^–3 P_i, the formation of proteoid roots did not affect plant dry matter yields or shoot to root dry matter ratios, indicating that proteoid roots can form under conditions of adequate P supply and not at the expense of dry matter production. Plants with over 50% of the root system as proteoid roots had tissue P concentrations considered adequate for maximum growth, providing additional evidence that proteoid roots can form on P-sufficient plants. There was an inverse relationship between the P_i concentration in the youngest mature leaf and proteoid root formation. Citrate efflux and the activities of enzymes associated with citric acid synthesis (phosphoenolpyruvate carboxylase and malate dehydrogenase) varied along the proteoid root axis, being greatest in young proteoid rootlets of the 1–3 cm region from the root tip. Citrate release from the 0–1 and 5–9 cm regions of the proteoid root was only 7% (per unit root length) of that from the 1–3 cm segment. Electrical potential and ³²P_i uptake measurements showed that P_i uptake was more uniform along the proteoid root than citrate efflux.