Higher plant diversity enhances soil stability in disturbed alpine ecosystems

Genotypic differences in acquiring immobile P exist among species or cultivars within one species. We investigated the P-efficiency mechanisms of rapeseed (Brassica napus L.) in low P soil by measuring plant growth, P acquisition and rhizosphere properties. Two genotypes with different P efficiencies were grown in a root-compartment experiment under low P (P15: 15 mg P kg^?1) and high P (P100: 100 mg P kg^?1) treatments. The P-efficient genotype produced more biomass, and had a high seed yield and high P acquisition efficiency under low P treatment. Under both P treatments, both genotypes decreased inorganic P (Pi) and organic P (Po) fractions in the rhizosphere soil. However there was no decrease in NaHCO₃-Po at P100. For the P15 treatment, the concentrations of NaHCO₃-Po and NaOH-Po were negatively correlated with soil acid phosphatase activity. The P-efficient genotype 102 differed from the P-inefficient genotype 105 in the following ways. In the rhizosphere the soil pH was lower, acid phosphatase activity was higher, and depletion of P was greater. Further the depletion zones were wider. These results suggested that improving P efficiency based on the character of P efficiency acquisition in P-efficient genotype would be a potential approach for maintaining rapeseed yield potential in soils with low P bioavailability.     

Effects of transgenic soybean on growth and phosphorus acquisition in mixed culture system

Transgenic soybean plants overexpressing the Arabidopsis purple acid phosphatase gene AtPAP15 (OXp) or the soybean expansin gene GmEXPB2 (OXe) can improve phosphorous (P) efficiency in pure culture by increasing Apase secretion or changing root morphology. In this study, soybean‐soybean mixed cultures were employed to illuminate P acquisition among plants in mixed stands of transgenic and wild‐type soybean. Our results showed that transgenic soybean plants were much more competitive, and had greater growth and P uptake than wild‐type soybean in mixed culture in both low P calcareous and acid soils. Furthermore, OXe plants had an advantage in calcareous soils when mixed with OXp, whereas the latter performed much better in acid soils. In soybean‐maize mixed culture, transgenic soybean had no impact on maize growth compared to controls in both acid and calcareous soils with different P conditions. As for soybean in mixed culture, OXp plants had no significant advantages regardless of P availability or soil type, while P efficiency improved in OXe in calcareous soils compared to controls. These results imply that physiological traits could be easily affected by the mixed maize. Transgenic soybean plants with enhanced root traits had more competitive advantages than those with improved root physiology in mixed culture.     

Phosphorus translocation in salt-stressed cotton

The effect of salinity on plants has usually been studied at high inorganic P concentration ([Pi]) in the nutrient solution, and salinity × Pi interactions have been examined at much higher [Pi] than found in soil solutions. Short-term ³²Pi experiments were carried out to study the effect of salinity (150 m M NaCl) on phosphorus translocation in cotton plants ( Gossypium hirsutum L. cv. Acala SJ-2) grown in nutrient solutions containing 10 μ M [Pi]. The effect of additional Ca to a concentration of 10 μ M was also tested. Salinity inhibited ³²P translocation from root to shoot. This inhibition was more evident at higher [Pi] in the root medium. Increasing [Pi] 33-fold in the solution resulted in a 4.3-fold increase in [³²P] in the root under saline conditions, but only in a 1,8-fold increase in the shoot. In older shoot tissues total [P] was elevated in the salinized plants. In the young tissues, however, total P concentration was higher in control plants. Inhibition of ³²P translocation by salinity was greater from root to young leaves than to mature shoot tissues. Salinity also decreased ³²P recirculation from the cotyledons to the young leaf. Inhibition by salinity of both ³²P translocation and recirculation to young leaves was fully reversed by increasing Ca supply from 1 to 10     

Growth response of komatsuna (Brassica rapa var. peruviridis) to root and foliar applications of phosphite

Soil and hydroponic culture experiments were conducted to investigate the effects of phosphite (Phi) as phosphorus (P) fertilizer via root and foliar applications on the growth and P supply of komatsuna. In both experiments, root P treatments were combinations of Phi and phosphate (Pi) at different Pi:Phi ratios, for a total of high P level (92 mg P pot^?1; the soil experiment) or low P level (0.05 mM P; the hydroponic experiment). Foliar P treatments were deionized water (control), a Pi solution and a Phi solution at low concentration of 0.05% P₂O₅. In both experiments, shoot dry weight of plants significantly decreased as Pi:Phi ratio decreased. In the soil experiment, plants grew abnormally at a Pi:Phi ratio of 25:75 and died when P was applied to soil entirely as Phi form (0:100 treatment). In the hydroponic experiment, no visible damage was found in shoot but root growth was strongly inhibited with severe damage symptoms at low Pi:Phi ratios. Total P concentration in plant decreased significantly with decreasing Pi:Phi ratio, especially in the hydroponic experiment. Foliar application of Phi although greatly increased total P of plants compared to that of Pi in both experiments, it did not improve but further decreased plant growth at low Pi:Phi ratios in the soil experiment and at all Pi:Phi ratios in the hydroponic experiment. The results of this study clearly indicated that Phi could not be used as P fertilizer by komatsuna plants via both application methods and could not substitute P at any rate at either low or high level. No beneficial effect of Phi was detected even when it was applied at low rate or applied in combination with Pi at different ratios. The effects of Phi were strongly dependent on the P nutrition status of plants; and plants that were not sufficiently fertilized with Pi may become vulnerable to Phi even at low levels.     

High external phosphate (Pi) increases sodium ion uptake and reduces salt tolerance of 'Pi-tolerant' soybean

Previous studies on the interaction between environmental inorganic phosphate (Pi) and salinity stress using soybean cultivars sensitive to high external Pi had two limitations: (1) the phenotype was dominated by overaccumulation of phosphorus (P); and (2) no detailed analysis was performed for sodium ion uptake. In this study, we focused on the effects of high external Pi on the sodium ion uptake in 'Pi-tolerant' soybean cultivars. The P accumulation in Pi-tolerant soybean Union was much lower [9.0 mg g⁻¹ dry weight (DW); contrasting to 38–76 mg g⁻¹ DW in the 'Pi-sensitive' soybean cultivars]. At in planta level, high level of external Pi significantly ( P  < 0.001) increased net sodium ion uptake and aggravated salinity stress symptoms. The effects of high external Pi diminished when de-rooted plants were used, suggesting that root is the primary organ interacting with Pi in the growth medium. Two-cell models, including soybean suspension cells and the tobacco Bright Yellow-2 cell line, were also employed to study the effects of high external Pi at the cellular level. Consistent to in planta results, high external Pi uplifted cellular sodium ion uptake and reduced cell viability under salinity stress. Gene expression analyses further showed that HPi (2 m M Pi supplements; excessive level of Pi) could reduce the fold of induction of GmSOS1 and GmCNGC under salinity stress, suggesting that they may be possible molecular targets involved in the interaction between high external Pi and Na⁺ uptake.     

Effects of root-knot nematodes on cucumber leaf N and P contents, soil pH, and soil enzyme activities

A pot experiment was conducted to study the effects of inoculation with root-knot nematodes on the cucumber leaf N and P contents, and the rhizospheric and non-rhizospheric soil pH and enzyme activities. The rhizospheric soil pH didn't have a significant decrease until the inoculation rate reached 6000 eggs per plant. With the increase of inoculation rate, the leaf N and P contents, rhizospheric soil peroxidase activity, and rhizospheric and non-rhizospheric soil polyphenol oxidase activity all decreased gradually, rhizospheric soil catalase activity was in adverse, non-rhizospheric soil pH decreased after an initial increase, and non-rhizospheric soil catalase activity had no regular change. After inoculation, rhizospheric soil urease activity decreased significantly, but rhizospheric and non-rhizospheric soil phosphatase activity and non-rhizospheric soil peroxidase activity only had a significant decrease under high inoculation rate. In most cases, there existed significant correlations between rhizospheric soil pH, enzyme activities, and leaf N and P contents; and in some cases, there existed significant correlations between non-rhizospheric soil pH, enzyme activities, and leaf N and P contents.     

模拟氮沉降对低磷胁迫下马尾松不同家系根系分泌和磷效率的影响

近年来大气氮(N)沉降的增加, 导致森林土壤中有效N含量增加、N:P发生改变, 研究N沉降对低磷(P)胁迫下林木根系分泌和P效率的影响具有重要意义。该文以马尾松(Pinus massoniana)家系作为试验材料, 设置模拟N沉降与同质低P (介质表层与深层均缺P)、异质低P (介质表层P丰富、深层缺P)耦合的二年生盆栽实验, 系统研究了模拟N沉降对低P胁迫下马尾松根系分泌性酸性磷酸酶(APase)活性、有机酸分泌以及P效率的影响。结果表明: (1)同质低P和异质低P下, 模拟N沉降均显著提高了植株N:P化学计量比、增加了P素的相对匮乏程度, 从而诱导根系增加了APase和有机酸的分泌, 而同质低P比异质低P下增加幅度更大, 其中有机酸分泌均与马尾松生长呈正相关关系, 而APase活性与P效率相关性较小; (2)同质低P下, N沉降虽然增加了根系分泌, 但未提高马尾松P素吸收和生长量, 其原因在于, 同质低P下植株N:P过高, 因而植株对N沉降敏感性低; 在异质低P下, 植株表现为N、P共同限制, 因而对N敏感性较高, N沉降增加了根系分泌, 同时提高了N和P吸收效率、增加了生物量; (3)马尾松根系分泌对模拟N沉降的响应存在较大的家系差异。同质低P下, 家系71×20的有机酸分泌和生物量对N沉降的响应幅度较大; 异质低P下, 家系36×29、71×20和73×23对N沉降的响应幅度较大。     

Comparative Study of the Interactive Effects of Salinity and Phosphorus Availability in Wild (Hordeum maritimum) and Cultivated Barley (H. vulgare)

The present study aimed to compare the effects of phosphorus (P) deficiency applied only or combined with salinity on root response, P partitioning, acid phosphatase activity, and phenolic compounds in wild (Hordeum maritimum) and cultivated (H. vulgare) barley species. Seedlings were grown hydroponically under low or sufficient P supply, with or without 100 mM NaCl for 55 days. Results showed that, when individually applied, P deficiency and salinity restricted the whole plant relative growth rate in both species of barley, with a more pronounced impact of the former stress. These depressive effects were more pronounced in H. vulgare than in H. maritimum. The combined effects of P deficiency and salinity were not additive neither on whole plant RGR nor on root response parameters in both species. The root area, root/shoot P content, root and leaf acid phosphatase activities, and shoot flavonoids contents increased under P deficiency conditions with and without salt in both species. Overall, the relatively better tolerance of H. maritimum plants to P deficiency applied only or combined with salinity could be explained by the capacity of this species to maintain higher P acquisition efficiency in concomitance with a larger root system, a higher root/shoot DW ratio, a higher root/shoot P content, a greater root and leaf acid phosphatase activities, and a higher flavonoid content and antioxidant capacity under combined effects of both stresses. Thus, H. maritimum constitutes a promising model to ameliorate the tolerance of the cultivated barley species under low-P soils and/or saline regions.     

Overexpression of β‐expansin gene GmEXPB2 improves phosphorus efficiency in soybean

Soybean (Glycine max) is an important oil crop in agricultural production, but low phosphorus (P) availability limits soybean growth and production. Expansin is a family of plant cell wall proteins and involved in a variety of physiological processes, including cell division and enlargement, root growth and leaf development. To test the potential effects of expansins on crop production, we have developed soybean transgenic plants overexpressing a soybean β‐expansin gene GmEXPB2, which was significantly induced by phosphate (Pi) starvation. The results indicated that constitutive overexpression of GmEXPB2 promoted leaf expansion, sequentially stimulated root growth and consequently resulted in improved P efficiency in the transgenic plants under P‐limited conditions in hydroponics. In particular, when tested in calcareous (CS) and acid soils (AS), the two GmEXPB2 transgenic soybean lines showed above 25 and 40% increases in plant dry weight and P content, respectively to wild‐type plants in low‐P CS, but not in AS. To our knowledge, this is the first report in which improvement of P efficiency could be achieved through constitutive overexpression of an endogenous EXPB gene in soybean. These findings suggest that genetic modification of root and leaf traits might be a suitable strategy for improving crop production in low‐P soils.     

Apoplastic accumulation of iron in the epidermis of maize (Zea mays) roots grown in calcareous soil

High concentrations of Fe in the roots of plants grown in calcareous soil have been found in a variety of plants, which, nevertheless, show Fe deficiency symptoms. In the present work, energy dispersive X-ray (EDX) analysis at the cellular level has been used to characterize high root Fe concentrations in maize ( Zea mays L.) grown in a calcareous soil in comparison with low root Fe concentrations under acidic soil conditions. Roots were thoroughly washed to remove adhering soil particles from the root surface as far as possible. To avoid any interference with possibly still present soil particles, the excitation beam was focused on radial walls of neighboring cells as well as on the symplast. Under alkaline conditions, high Fe concentrations in the m M range and higher accumulated in the epidermal root apoplast. Symplastic Fe was not detectable. Only traces of Fe were detectable in the apoplast of the cortex parenchyma. Under acidic conditions, apoplastic root Fe concentrations were clearly lower than under alkaline conditions, and no Fe was detectable in the root apoplast by use of EDX analysis. We conclude that, under alkaline conditions, high amounts of Fe are trapped in the epidermal root apoplast (apoplastic Fe inactivation), probably because of a high apoplastic pH and thus restricted translocation towards the root stele and to the upper plant parts. In contrast, on acidic soils Fe translocation towards the root stele and thus Fe supply to the upper plant parts was not impaired. Our findings imply that Fe deficiency on calcareous soils is not caused by restricted acquisition of Fe from the soil.     

Preferential utilization of organic and inorganic sources of phosphorus by wheat plant

On soils of low P supply organic P (Po) makes up a similar or even larger part in soil solution than inorganic P (Pi). The ability of wheat (Triticum aestivum L., cv. Star) plants to hydrolyze and absorb this Po in comparison to similar concentrations of Pi was studied. Four concentration levels of Pi and Po were obtained by extracting two soils with deionized water in a ratio of 1:1 and concentrating the resulting filtrate by freeze drying to different degrees. The concentration of Pi varied between 5 and 36 μM and Po between 3 and 22 μM. Wheat seedlings were grown in these solutions for 12 and 24 h and acid and alkaline phosphatase activity determined. The reduction of Po concentration in solution expressed on a root length basis gave the rate of Po hydrolysis and the reduction in concentration of Pi and Po gave the P inflow into the roots. No alkaline phosphatase activity was detected. The activity of wheat root acid phosphatase increased with Po concentration in solution. Phosphorus uptake was 2 to 6 fold higher from Pi than from Po at similar concentrations of both. The rate of uptake from Pi, the inflow, as well as the rate of hydrolysis of Po increased linearly with concentration but at similar concentration the inflow was 2 to 4 times higher than the rate of Po hydrolysis. Results suggest that plants can utilize Po after hydrolysis by phosphatase, but Pi is more important and preferentially used by plants; Po may be essential for plant nutrition especially in high P-fixing soils.     

Response of root and sediment phosphatase activity to increased nutrients and salinity

Wetlands of northern Belize, distributed along a salinity gradient, are strongly phosphorus limited and dominated largely by three species of emergent macrophytes: Eleocharis cellulosa, Cladium jamaicense, and Typha domingensis. We assessed changes in root and sediment phosphatase activities of each species to simultaneous changes of nutrients (N, P) and salinity in a mesocosm experiment. Phosphorus and nitrogen treatment effects on both root and sediment phosphatase were highly significant for all the species, while salinity significantly affected root phosphatase activity in Cladium only. All species showed a significant negative correlation between root phosphatase activity and increasing tissue P content until a threshold of 0.2% P, 0.15% P and 0.12% P in Eleocharis, Cladium and Typha, respectively. There was also a significant negative correlation between soil available P and root and sediment phosphatases in all species. Activity of root phosphatases of Eleocharis and Typha were positively correlated with root tissue N. Both root and sediment phosphatases of all three species were positively correlated with soil available N. The strongest (positive) correlation was found between phoshatase activites and N/P ratios. The results confirmed that these systems are P-limited and that extracellular phosphatases respond to P enrichment by decreasing their activities. Expression of root phosphatase activity by dry root weight, sediment volume, or whole plant biomass gave very different relative results across nutrient treatments and species, suggesting that root phosphatase activities need to be interpreted in a wider context that considers root density.     

Soil solution phosphate,root uptake kinetics and nutrient acquisition: implications for a patchy soil environment

Summary The importance of increased root phosphate (P) uptake kinetics, root proliferation and local increases of soil solution P (P₁) for P acquisition from fertile soil microsites was explored with a simulation model and calculated uptake was compared with experimental data. Based on the partitioning of added P in microsites to P₁ and P adsorbed on soil particles and the results of a dual-isotope-labeling experiment (Caldwell et al. 1991a), acquisition of P from the fertile microsites was some 20 X that of uptake from an equal volume of soil which received only water. Simulations were in general agreement and also showed that elevation of root P uptake kinetics could contribute more to the increased acquisition than did root proliferation under these circumstances. Although increased physiological uptake capacity for P has generally been considered to be of little benefit because of diffusion limitation, in patchy soil environments selective elevation of P uptake kinetics in fertile microsites may be of considerable benefit. These tests were conducted in calcareous soil which releases much less P into the soil solution than do many other soils. In many noncalcareous soils the benefits of selective elevation of root uptake kinetics would likely be greater.     

Salt tolerance in Aster tripolium L. III. Na and K fluxes in intact seedlings

Abstract Uptake and transport of Na and K was studied using the radioactive tracers ²²Na and ⁴²K in intact Aster tripolium L. seedlings grown at two salinities CS 10 and CS 100, (containing 10mol m^?1 and 100 mol m^?3 Na, respectively, together with other major ions in the proportions found in sea water). At both salinities a much greater proportion of the Na than K taken up by the plant was subsequently transported to the shoot. Most ⁴²K fluxes were reduced by about 40% in CS 100 plants relative to CS 10 except root accumulation which increased. Experiments involving changing the salinity from CS 10 to CS 100 showed that ⁴²K fluxes remained constant for at least 40 h, indicating that competition with Na for uptake sites was not the cause of the reduced flux in CS 100 plants. ²²Na fluxes responded immediately to a change in salinity with all fluxes increasing six-fold when the salinity was raised. When the salinity was lowered, however, root accumulation returned to the level in CS 10 control plants whereas transport to the shoot was inhibited by the previous high salinity treatment, being reduced to only 35% of the rate in CS 10 plants. The time courses of osmotic adjustment and Na accumulation following an increase in salinity were found to be very similar, with sufficient Na being accumulated to account for the observed increase in sap osmotic pressure.     

Root surface phosphatase activity in ecotypes of Aegilops peregrina

The relationships between root surface phosphatase activity and the edaphic factors of their native habitats were investigated in four ecotypes of Aegilops peregrina (Hack.) Maire et Weil. In one set of experiments plants were grown in phosphate-deficient nutrient solution cultures (5 μ M ) with three pH values: 5.5, 6.5 and 7.5. In a second series, plants were grown in both P-poor and P-rich soils.
Results showed an optimal activity of the commonly-described root surface acid phosphatase of pH 4.5–5.0 in the ecotypes Meron (a P-poor montmorillonitic, typical mediterranean Terra-Rossa soil) and Har-Hurshan (a P-rich calcareous soil). However, in the ecotypes Malkiya (a P-rich kaolinitic Terra-Rossa) and Bet-Guvrin (a P-rich calcareous soil) the optimal activity of the phosphatase occurred at pH 6.0. The pH level of the growth solution had no effect on the pH of optimal activity of the phosphatase in the ecotypes Malkiya and Bet-Guvrin, but it somewhat affected their level of activity.
Phosphatase activity was stimulated when plant roots were grown in a P-poor soil, as compared to the activity of those which were grown in a P-rich soil. Plants of the Malkiya ecotype exhibited the strongest activation of phosphatase as compared to the other three ecotypes. It seems that ecotypes which have evolved in P-rich soils may regulate their root surface phosphatase activity better than those which have evolved in P-poor soils.     

NaCl‐induced accumulation of glycinebetaine in four subtropical halophytes from Pakistan

Variation in both the total content and the concentration of glycinebetaine in response to increasing soil salinity was studied in the salt‐secreting Atriplex griffithii Moq., the leaf succulent Suaeda fruticosa (L.) Forssk., the stem succulent Haloxylon recurvum Bunge ex Boiss. and the osmotically adjusting desert grass Halopyrum mucronatum (L.) Stapf. collected from a subtropical maritime desert in Pakistan. Glycinebetaine content (mmol kg⁻¹ dry weight) increased with increasing NaCl in Atriplex griffithii, Haloxylon recurvum and Halopyrum mucronatum , but peaked at 600 m M NaCl for Suaeda fruticosa and declined thereafter. Glycinebetaine concentration (mmol l⁻¹ tissue water) increased with increasing salinity in all four halophyte species and was sufficiently high to serve as an osmoticum in all cases.     

Response of wheat to subsoil salinity and temporary water stress at different stages of the reproductive phase

To examine the effects of subsoil NaCl salinity in relation to water stress imposed at different growth stages, wheat was grown in a heavy texture clay soil (vertosol) under glasshouse conditions in polythene lined cylindrical PVC pots (100 cm long with 10.5 cm diameter) with very low salinity level (EC_e 1.0 dS/m; ESP 1.0 and Cl 30 mg/kg soil) in top 10 cm soil (10–20 cm pot zone) and low salinity level (EC_e 2.5 dS/m, ESP 5, and Cl 100 mg/kg soil) in top 10–20 cm soil (20–30 cm pot zone). The plants were exposed to three subsoil salinity levels in the 20–90 cm subsoil (30–100 cm pot zone) namely low salinity (EC_e: 2.5 dS/m, ESP: 5, Cl: 100 mg/kg soil), medium salinity (EC_e: 4.0 dS/m, ESP: 10, Cl: 400 mg/kg) and high salinity (EC_e: 11.5 dS/m, ESP: 20, Cl: 1950 mg/kg) in the subsoil (20–90 cm soil layer: 30–100 cm pot zone). Watering of plants was withheld for 20 days commencing at either early booting or anthesis or mid grain filling, and then resumed until maturity, and these treatments were compared with no water stress. Water stress commencing at anthesis stage had the most depressing effect on grain yield and water use efficiency of wheat followed by water stress at grain filling stage and early booting stage. High subsoil salinity reduced grain yield by 39.1, 24.3%, and 13.4% respectively in plants water-stressed around anthesis, early booting, and mid grain filling compared with 36.6% in well-watered plants. There was a significant reduction in root biomass, rooting depth, water uptake and water use efficiency of wheat with increasing subsoil salinity irrespective of water regimes. Plants at high subsoil salinity had 64% of their root biomass in the top 0–30 cm soil and there was a marked reduction in subsoil water uptake. Roots also penetrated below the non-saline surface into salinised subsoil and led to attain high concentration of Na and Cl and reduced Ca/Na and K/Na ratio of flag leaf at anthesis stage. Results suggest that high subsoil salinity affects root growth and water uptake, grain yield and water use efficiency even in well water plants. Water stress at anthesis stage had the most depressing effect on wheat.     

Soluble inorganic tissue phosphorus and calcicole-calcifuge behaviour of plants

BACKGROUND AND AIMS: Natural and semi-natural, non-fertilized calcareous soils are consistently low in soluble and easily exchangeable phosphate. An over-utilization, or possibly an immobilization, of inorganic P in the tissues of calcifuge plants may take place, if such plants are forced to grow on a calcareous soil, though this has not been experimentally demonstrated. The objectives of this study are, therefore, to elucidate if calcifuge plants, when forced to develop on a calcareous soil, not only have lower total P (Ptot) concentrations in their leaves than calcicole plants grown on such soil, but also a lower proportion of Ptot as water-soluble, inorganic phosphate. Such differences may be of importance in understanding the calcicole-calcifuge behaviour of plants. MATERIALS AND METHODS: Plants of five calcicole and five calcifuge herbs and three calcicole and three calcifuge grasses were cultivated in a glasshouse on a moderately acid Cambisol and a calcareous Rendzic Leptosol using seeds of wild populations from southern Sweden. The calcifuges were: Corynephorus canescens, Deschampsia flexuosa, Holcus mollis, Digitalis purpurea, Lychnis viscaria, Rumex acetosella, Scleranthus annuus and Silene rupestris. The calcicoles were: Melica ciliata, Phleum phleoides, Sesleria caerulea, Arabis hirsuta, Sanguisorba minor, Scabiosa columbaria, Silene uniflora ssp. petraea and Veronica spicata. KEY RESULTS: At harvest, calcifuges had much lower leaf tissue concentrations of Ptot and Pi than calcicoles when grown on the calcareous soil, and biomass production of the calcifuges was poor on this soil. Moreover, the calcifuge herbs had, on average, a lower proportion of their Ptot as Pi than had the calcicole herbs. The calcifuge herbs were also unable to avoid excessive uptake of Ca from the calcareous soil. The calcifuge grasses maintained a similar proportion of Ptot as Pi as the calcicole grasses, but their growth was still poor on the calcareous soil. CONCLUSIONS: On calcareous soil, very little Pi in the tissues of calcifuge herbs is, at any time, available for use in various physiological functions. This is of importance to their photosynthesis, growth, competition and final survival on such soils.