Occurrence of arbuscular mycorrhizal fungi in a phosphorus-poor wetland and mycorrhizal response to phosphorus fertilization

The presence of arbuscular mycorrhizas in fens has received little attention, but because fen plants are often phosphorus limited, the plant-fungus interaction could be an important factor in plant competition for phosphorus. In this field study, we determined mycorrhizal colonization rates for 18 fen plant species. Also in the field, we examined the effect of four different forms of phosphorus on the percentage colonization for one fen plant species, Solidago patula. We found that in a species-rich, phosphorus-poor wetland both mycorrhizal and nonmycorrhizal species were common. Nine of ten dicotyledonous species examined formed arbuscular mycorrhizas, while all monocotyledonous species were at most very weakly mycorrhizal. A morphological explanation for this pattern is that the monocots in our study have more extensive aerenchyma, especially in coarse roots. Therefore, monocots are able to transport oxygen to their roots more effectively than dicots. In the organic wetland soil, additional oxygen in the rhizosphere promotes phosphorus mineralization and availability. Two of the monocot species (Typha latifolia and Carex lasiocarpa), which have been described previously as mycorrhizal in other wetland types, are surprisingly nonmycorrhizal in our phosphorus-poor study site, suggesting that a mycorrhizal association would not offer improved phosphorus nutrition to these species. In contrast, our field phosphorus addition decreased mycorrhizal colonization in S. patula, suggesting that one benefit to S. patula of the mycorrhizas is phosphorus uptake.     

Effect of phosphorus fertilization on water stress in Douglas fir seedlings during soil drying

A growth chamber experiment was conducted to determine if P fertilization to enhance the P nutrition of otherwise N and P deficient Douglas fir [Pseudotsuga menziesii (Mirb.) Franco] seedlings reduces water stress in the seedlings during drought periods. Seedlings were grown in pasteurized mineral soil under well-watered conditions and fertilized periodically with a small amount of nutrient solution containing P at either of three levels: 0, 20, or 50 mg P L^-1. By age 6 mo, leaf nutrient analysis indicated that N and P were deficient in control (0 mg P L^-1) seedlings. The highest level of P fertilization, which doubled leaf P concentration, did not affect plant biomass, suggesting that N deficiency was limiting growth. When these seedlings were subjected to drought, there was no effect of P fertilization on leaf water potential or osmotic potential. Furthermore, P fertilized seedlings had lower stomatal conductance and net photosynthesis rate. These results indicate that enhanced P nutrition, in the presence of N deficiency, does not reduce water stress in Douglas fir seedlings during drought periods.     

Leaf water relations and maintenance of gas exchange in coffee cultivars grown in drying soil

Plant water status, leaf tissue pressure-volume relationships, and photosynthetic gas exchange were monitored in five coffee (Coffea arabica L.) cultivars growing in drying soil in the field. There were large differences among cultivars in the rates at which leaf water potential (Ψ_L) and gas exchange activity declined when irrigation was discontinued. Pressure-volume curve analysis indicated that increased leaf water deficits in droughted plants led to reductions in bulk leaf elasticity, osmotic potential, and in the Ψ_L at which turgor loss occurred. Adjustments in Ψ_L at zero turgor were not sufficient to prevent loss or near loss of turgor in three of five cultivars at the lowest values of midday Ψ_L attained. Maintenance of protoplasmic volume was more pronounced than maintenance of turgor as soil drying progressed. Changes in assimilation and stomatal conductance were largely independent of changes in bulk leaf turgor, but were associated with changes in relative symplast volume. It is suggested that osmotic and elastic adjustment contributed to maintenance of gas exchange in droughted coffee leaves probably through their effects on symplast volume rather than turgor.     

Leaf water relations and solute accumulation in two grain sorghum lines exhibiting contrasting drought tolerance

A drought-tolerant grain sorghum line (K886) maintained significantlyhigher relative water content (RWC), osmotic potential at fullturgor (     

Nodulation and growth response ofAllocasuarina andCasuarina species to phosphorus fertilization

To examine how soil phosphorus status affects nitrogen fixation by the Casuarinaceae —Frankia symbiosis,Casuarina equisetifolia and two species ofAllocasuarina (A. torulosa andA. littoralis) inoculated or fertilized with KNO₃ were grown in pots in an acid soil at 4 soil phosphate levels. InoculatedC. equisetifolia nodulated well by 12 weeks after planting and the numbers and weight of nodules increased markedly with phosphorus addition. Growth ofC. equisetifolia dependent on symbiotically fixed nitrogen was more sensitive to low levels of phosphorus (30 mg kg^–1 soil) than was growth of seedings supplied with combined nitrogen; at higher levels of phosphorus, the growth response curves were similar for both nitrogen fertilized and inoculated plants. The interaction between phosphorus and nitrogen treatments (inoculated and nitrogen fertilized) demonstrated that there was a greater requirement of phosphorus for symbiotic nitrogen fixation than for plant growth when soil phosphorus was low.WithAllocasuarina species, large plant to plant variation in nodulation occurred both within pots and between replicates. This result suggests genetic variation in nodulation withinAllocasuarina species. Nodulation ofAllocasuarina species did not start until 16 weeks after planting and no growth response due toFrankia inoculation was obtained at the time of harvest. Addition of nitrogen starter is suggested to boost plant growth before the establishment of the symbiosis. Growth ofAllocasuarina species fertilized with nitrogen responded to increasing levels of phosphorus up to 90 mg P/kg soil after which it declined by 69% forA. littoralis. The decrease in shoot weight ofA. littoralis, A. torulosa, C. equisetifolia andC. cunninghamiana at high phosphorus was confirmed in a sand culture experiment, and may be atributable to phosphorus toxicity.     

Leaf traits of two Mediterranean perennial tussock grass species in relation to soil nitrogen and phosphorus availability

Studying relationships of plant traits to ecosystem properties is an emerging approach aiming to understand plant's potential effect on ecosystem functioning. In the current study, we explored links between morphological and nutritional leaf traits of two Mediterranean perennial grass species Stipa tenacissima and Lygeum spartum, widely used to prevent desertification process by stabilizing sand dunes. We evaluated also relationships in terms of nitrogen (N) and phosphorus (P) availability between leaves of the investigated species and the corresponding soil. Our results showed that leaf P was very low in comparison with leaf N for the two investigated species. In fact, chlorophyll content, photosynthesis capacity and water conservation during photosynthesis are mainly linked to leaf nitrogen content. Our findings support previous studies showing that at the species levels, morphological and nutritional leaf traits were not related. On the other hand, significant relationships were obtained between soil N and leaf N for S. tenacissima (P = 0.011) and L. spartum (P = 0.033). However, leaf P was not significantly related to soil P availability for both species. We suggest that any decrease in soil N with the predicted increasing aridity may result in reduction in leaf N and thus in worst dysfunction of some biological processes levels.     

Responses of leaf gas exchange, water relations, and water consumption in seedlings of four semiarid tree species to soil drying

To understand the response patterns to soil drying and the water use properties of commonly reforested trees in the semiarid Loess Plateau region of China, a glasshouse experiment was carried out with the seedlings of four species, i.e., Robinia pseudoacacia, Armeniaca sibirica, Syringa oblata, and Quercus liaotungensis. Severe water stress induced by withholding water resulted in permanent wilting of most of the seedlings pot-cultured with sandy soil in 8–12 days. Predawn and midday leaf water potentials and gas exchange characteristics (e.g., stomatal conductance) in the seedlings did not show marked changes until the volumetric soil water content decreased to about 0.05. As the soil water content decreased further, these physiological parameters rapidly declined, approaching their minimal levels at the stage of permanent wilting. The response of each parameter to soil water content changes was fitted with a non-linear saturation curve. Though the results suggested that the general pattern of responses to soil drying was identical among the species, quantitative differences in drought tolerance and water use properties were detected. Leaf stomatal conductance in R. pseudoacacia and A. sibirica showed earlier responses to reduced predawn leaf water potentials. However, water use characteristics and specific leaf area indicated that these two species consumed more water and may not be as drought tolerant as S. oblata and Q. liaotungensis. These results may provide important information to compare the reforestation species with respect to soil drying.     

Biomass partition,leaf gas exchange and water relations of alfalfa and milkvetch seedlings in response to soil drying

This study compared physiological and growth responses to water stress of two legume species during the seedling stage. Potted alfalfa (Medicago sativa L. cv. Algonquin) and milkvetch (Astragalus adsurgens Pall. cv. Pengyang earlymaturing vetch) seedlings were grown under well-watered [soil water content (SWC) maintained at 14.92% daily] or water-stressed conditions (drying) for 15 days. Net photosynthetic rate (P _N), transpiration rate (E) and stomatal conductance (g _s) of both species decreased parabolically. When SWC decreased to 7.2% and 10.3%, g _s values for alfalfa and milkvetch were significantly different from those of the respective well-watered plants (p<0.05). When SWC decreased to 6.6% for alfalfa and 6.8% for milkvetch, leaf water potentials (ψ_L) were significantly different from those of the well-watered plants (p<0.05). Thus the difference between the SWC thresholds for a nonhydraulic root signal (nHRS) and a hydraulic root signal (HRS) were 0.6% and 3.5% for alfalfa and milkvetch, respectively. Milkvetch had a lower g _s than alfalfa for a given SWC (p<0.05). Although alfalfa seedlings had a higher dry mass (DM) and root:shoot ratio (R/S) than milkvetch in both treatments (p<0.05), we concluded that milkvetch seedlings had greater drought tolerance than alfalfa.     

Plant water relations and control of cell elongation at low water potentials

Recent developments in water status measurement techniques using the psychrometer, the pressure probe, the osmometer and pressure chamber are reviewed, and the process of cell elongation from the viewpoint of plant-water relations is discussed for plants subjected to various environmental stress conditions. Under water-deficient conditions, cell elongation of higher plants can be inhibited by interruption of water flow from the xylem to the surrounding elongating cells. The process of growth inhibition at low water potentials could be reversed by increasing the xylem water potential by means of pressure application in the root region, allowing water to flow from the xylem to the surrounding cells. This finding confirmed that a water potential field associated with growth process,i.e., the growth-induced water potential, is an important regulating factor for cell elongation other than metabolic factors. The concept of the growth-induced water potential was found to be applicable for growth retardation caused by cold stress, heat stress, nutrient deficiency and salinity stress conditions. In the present review, the fact that the cell elongation rate is primarily associated with how much water can be absorbed by elongating cells under water-deficiency, nutrient deficiency, salt stress, cold stress and heat stress conditions is suggested.     

Allocation trade-off between root and mycorrhizal surface defines nitrogen and phosphorus relations in 13 grassland species

Plant and Soil - In oligotrophic ecosystems efficient nutrient uptake mechanisms, like extensive root systems or the association with belowground symbionts (e.g. arbuscular mycorrhizal fungi, AMF),...     

Nitrogen limitation impairs plant control over the arbuscular mycorrhizal symbiosis in response to phosphorus and shading in two European sand dune species

The symbiosis of plants with arbuscular mycorrhizal fungi (AMF) may become parasitic if the cost:benefit ratio (carbon:phosphorus ratio) increases. In case of mycorrhizal parasitism, a plant may prevent growth depression through the reduction of root colonization as a form of control over the symbiosis. In this greenhouse study, we attempted to manipulate the cost:benefit ratio of the arbuscular mycorrhizal symbiosis by shading and/or phosphorus (P) fertilization in the differentially mycotrophic plant species Hieracium pilosella and Corynephorus canescens. By repeated sampling of soil cores, we assessed the temporal progress of plant investment towards mycorrhizal structures as a measure of plant control over the AMF. Unexpectedly, we found no obvious treatment effects on mycorrhizal growth dependency (MGD), most likely caused by constant N-limitation in AM plants being enhanced by P-fertilization and shade probably not exacerbating plant C-budget for AMF. This highlights the importance of N:P:C stoichiometry for the outcome of the symbiosis. Nevertheless, we found possible control mechanisms in shaded H. pilosella, with considerably higher resource investments into root than into hyphal growth, while root colonization was only marginally suppressed. This control only manifested after 4 weeks of growth under potentially detrimental conditions, emphasizing the importance of time in plant control over the arbuscular mycorrhizal symbiosis. In contrast, the less mycotrophic C. canescens did not exhibit obvious changes in mycorrhizal investments in reaction to shading and P-fertilization, possibly because the low mycotrophy and AMF colonization already imposes a functioning control mechanism in this species. Our study suggests that highly mycotrophic plants may have a stronger need to keep AMF in check than less mycotrophic plants, which may have implications for the role of mycotrophy in the outcome of symbiotic interactions in natural situations.     

Water relations and hydraulic control of stomatal behaviour in bell pepper plant in partial soil drying

Two experiments, a split-root experiment and a root pressurizing experiment, were performed to test whether hydraulic signalling of soil drying plays a dominant role in controlling stomatal closure in herbaceous bell pepper plants. In the split-root experiment, when both root parts were dried, synchronous decreases in stomatal conductance (g_s), leaf water potential (LWP) and stem sap flow (SF_stem) were observed. The value of g_s was found to be closely related to soil water potential (SWP) in both compartments. Tight relationships were observed between g_s and stem sap flow under all conditions of water stress, indicating a complete stomatal adjustment of transpiration. When the half-root system has been dried to the extent that its water uptake dropped to almost zero, declines in g_s of less than 20% were observed without obvious changes in LWP. The reduced plant hydraulic conductance resulting from decreased sap flow and unchanged LWP may be a hydraulic signal controlling stomatal closure; the results of root pressurizing supported this hypothesis. Both LWP and g_s in water-stressed plants recovered completely within 25 min of the application of root pressurizing, and decreased significantly within 40 min after pressure release, indicating the hydraulic control of stomatal closure. Our results are in contrast to those of other studies on other herbaceous species, which suggested that chemical messengers from the roots bring about stomatal closure when plants are in water stress.     

The response of two Glomus mycorrhizal fungi and a fine endophyte to elevated atmospheric CO2, soil warming and drought

Plantago lanceolata plants were grown under various environmental conditions in association with the mycorrhizal fungi Glomus mosseae, G. caledonium and a fine endophyte either individually or all together. Using a time‐course approach, we investigated the effects of elevated atmospheric CO₂ (eCO₂), soil warming and drought and their interactions on root length colonized (RLC) by mycorrhizal fungi and extraradical mycorrhizal hyphal (EMH) production. Plant growth responded as would be expected to the environmental manipulations. There was no plant growth‐independent effect of eCO₂ on mycorrhizal colonization; however, EMH production was stimulated by eCO₂, i.e. there was increased partitioning of below‐ground carbon to the EMH. Soil warming directly stimulated both percent RLC by the Glomus species and EMH density; soil warming did not affect RLC by the fine endophyte. Drought decreased percent RLC for the fine endophyte, but not for the Glomus species. The presence of one mycorrhizal fungus did not affect the response of another to the environmental variables. There was no evidence of any interactive effects of the environmental variables on RLC, but there were significant environmental interactions on EMH production. In particular, the stimulatory effects of eCO₂ and soil warming on EMH density were not additive. The results are discussed in terms of the soil carbon cycle, highlighting some crucial gaps in our knowledge. If future environmental changes affect mycorrhizal fungal turnover and respiration, then this could have important implications for the terrestrial carbon cycle.