Water use patterns of four co-occurring chaparral shrubs

Summary Mixed stands of chaparral in California usually contain several species of shrubs growing close to each other so that aerial branches and subterranean roots overlap. There is some evidence that roots are stratified relative to depth. It may be that root stratification promotes sharing of soil moisture resources. We examined this possibility by comparing seasonal water use patterns in a mixed stand of chaparral dominated by four species of shrubs: Quercus durata, Heteromeles arbutifolia, Adenostoma fasciculatum, and Rhamnus californica. We used a neutron probe and soil phychrometers to follow seasonal depletion and recharging of soil moisture and compared these patterns to seasonal patterns of predawn water potentials, diurnal leaf conductances, and diurnal leaf water potentials. Our results indicated that 1) Quercus was deeply rooted, having high water potentials and high leaf conductances throughout the summer drought period, 2) Heteromeles/Adenostoma were intermediate in rooting depth, water potentials, and leaf conductances, and 3) Rhamnus was shallow rooted, having the lowest water potentials and leaf conductances. During the peak of the drought, predawn water potentials for Quercus corresponded to soil water potentials at or below a depth of 2 m, predawn water potentials of Heteromeles/ Adenostoma corresponded to a depth of 0.75 m, and predawn water potentials of Rhamnus corresponded to a depth of 0.5 m. This study supports the concept that co-occurring shrubs of chaparral in California utilize a different base of soil moisture resources.     

Seasonal variation and annual fluctuation in granite outcrop plant communities

Permanent quadrats in granite outcrop plant communities allowed us to monitor seasonal variation and annual fluctuation in community structure. Seasonal species turn-over was significant in communities on shallow soil, but not in communities on deeper soil where seasonal dominance shifts were common. Exceptional meteorological events appeared to mediate phenomena of competitive release in some island communities. A decrease in the abundance of Arenaria uniflora in Lichen-annual island communities, following a spring drought, was correlated with an increase in the abundance of Sedum smallii, a shallower-soil species. Richness in Annual-perennial island communities was higher in spring 1985 than in 1984 or 1986, and this occurred as the dominant species, Senecio tomentosus, temporarily declined in importance following a severe drought in late summer 1984. Significant annual fluctuation in the cover of Viguiera porteri could also be related to variations in the summer precipitation regime. Overall, plant responses to drought were individualistic and depended largely on the timing of these meteorological events in relation to the life-stages and/or the physiological status of the plants.Abbreviations AP = Annual-perennial island communities - HST = Herb-shrub-tree island communities - LA = Lichen-annual island communities - SS = Sedum smallii island communities     

Root Distribution and Nutrient Status of Mycorrhizal and Non-mycorrhizal Pinus sylvestris L. Seedlings Growing in a Sandy Substrate with Lignite Fragments

Reclaimed mine soils of the Lusatian mining district are characterised by small-scale heterogeneous distribution of lignite fragments of varying size embedded in a matrix of Tertiary and Quaternary sandy material. Despite amelioration with basic fly ashes, ongoing pyrite oxidation and the subsequent acidification generate a high physical and chemical heterogeneity within the substrate, which could negatively affect root proliferation. We hypothesised that this limitation for the root system may be compensated for by intensive exploration of the porous lignite fragments by roots and/or mycorrhizal hyphae to access water and nutrients stored in these fragments. To test this hypothesis, we compared growth, shoot nutrient content, and root distribution of mycorrhizal and non-mycorrhizal Pinus sylvestris L. seedlings in lignite-containing and lignite-free sandy substrate. Rhizotrons used for this experiment were filled with a sandy matrix with 6–9 evenly distributed spots of lignite fragments. Treatments included different levels of water and nutrient availability. After 8 months of growth, root tip vitality as well as growth and shoot nutrient concentration of the plants was higher for treatments with lignite spots in the sandy substrate than for sandy substrate without such amendments. Compared to the non-mycorrhizal plants, the seedlings inoculated with Paxillus involutus (Batsch) Fr. had a higher root dry mass, an increased number of root tips and a higher root length. These results confirm our hypothesis that the lignite fragments are an important nutrient and water reservoir for plants in these mine soils and they indicate that mycorrhizal colonisation may allow an intensive exploration of porous lignite fragments by mycorrhizal hyphae.     

Quantification of water uptake by arbuscular mycorrhizal hyphae and its significance for leaf growth, water relations, and gas exchange of barley subjected to drought stress

Arbuscular mycorrhizal fungi alleviate drought stress in their host plants via the direct uptake and transfer of water and nutrients through the fungal hyphae to the host plants. To quantify the contribution of the hyphae to plant water uptake, a new split-root hyphae system was designed and employed on barley grown in loamy soil inoculated with Glomus intraradices under well-watered and drought conditions in a growth chamber with a 14-h light period and a constant temperature (15 degrees C; day/night). Drought conditions were initiated 21 days after sowing, with a total of eight 7-day drying cycles applied. Leaf water relations, net photosynthesis rates, and stomatal conductance were measured at the end of each drying cycle. Plants were harvested 90 days after sowing. Compared to the control treatment, the leaf elongation rate and the dry weight of the shoots and roots were reduced in all plants under drought conditions. However, drought resistance was comparatively increased in the mycorrhizal host plants, which suffered smaller decreases in leaf elongation, net photosynthetic rate, stomatal conductance, and turgor pressure compared to the non-mycorrhizal plants. Quantification of the contribution of the arbuscular mycorrhizal hyphae to root water uptake showed that, compared to the non-mycorrhizal treatment, 4 % of water in the hyphal compartment was transferred to the root compartment through the arbuscular mycorrhizal hyphae under drought conditions. This indicates that there is indeed transport of water by the arbuscular mycorrhizal hyphae under drought conditions. Although only a small amount of water transport from the hyphal compartment was detected, the much higher hyphal density found in the root compartment than in the hyphal compartment suggests that a larger amount of water uptake by the arbuscular mycorrhizal hyphae may occur in the root compartment.     

Comparative physiology of burned and unburned Rhus laurina after chaparral wildfire

Summary Laurel Sumac (Rhus laurina) is a dominant member of the coastal chaparral community of southern California that survives periodic burning by wildfires by resprouting from a lignotuber (root crown). We investigated the physiological basis for resprouting by comparing shoot elongation, leaf nitrogen content, tissue water status, leaf conductance to water vapor diffusion, and photosynthetic rates of post-fire R. laurina to those of adjacent unburned shrubs. Resprouts had higher rates of shoot elongation, leaf conductance, and photosynthesis than mature, unburned shrubs. Leaf nitrogen contents were elevated in burned shrubs even though their leaves developed interveinal chlorosis. A comparison of soil water potential to predawn water potential indicated that roots of R. laurina remain active below 2 m during the first summer drought after wildfire. Our results support the hypothesis that lignotubers not only contain dormant buds that develop into aerial shoots after wildfire but they also supply nutrient resources that enhance shoot elongation. Because R. laurina is relatively sensitive to drought, yet very successful in its rapid recovery after fire, maintaining an active root system after shoot removal may be the primary function of the massive lignotuber formed by this species.     

Superoxide dismutase and total peroxidase activities in relation to drought recovery performance of mycorrhizal shrub seedlings grown in an amended semiarid soil

We studied the effect of inoculation with a mixture of three arbuscular mycorrhizal (AM) fungi (Glomus intraradices Schenck & Smith, Glomus deserticola (Trappe, Bloss. & Menge) and Glomus mosseae (Nicol & Gerd.) Gerd. & Trappe) and addition of a composted organic residue on plant growth, nutrient uptake, mycorrhizal colonisation and superoxide dismutase (SOD, EC 1.15.1.1) and total peroxidase (POX, EC 1.11.1.7) activities in shoots of Juniperus oxycedrus seedlings after well-watered, drought and recovery periods. The mycorrhizal inoculation and composted residue addition significantly increased the growth, foliar nutrients (N, P, K) and shoot water content of the plants, independent of the water regime. POX activity in control plants increased during drought (about 250% higher than under well-watered conditions) and returned to initial levels after re-watering. The seedlings inoculated with AM fungi showed the highest values of POX activity, followed by the plants grown in the amended soil, which varied little during the drought and recovery periods. Drought decreased the SOD activity in shoots of both J. oxycedrus seedlings inoculated with AM fungi and those grown with composted residue, but did not affect that of control plants. After re-watering, the SOD activity in mycorrhizal or residue-amended plants increased, showing values similar to control plants.     

Endomycorrhizal fungal species mediate 15N transfer from soybean to maize in non-fumigated soil

The effect of mycorrhizal inoculation on ¹⁵N transfer from soybean to maize was studied in fumigated and non-fumigated soil. Three Glomus species and a non-inoculated control were compared.In spite of higher levels of root colonization and more abundant hyphae associated with plants growing in fumigated soil, mycorrhizae-enhanced ¹⁵N transfer to maize was significant only in non-fumigated plots. High ¹⁵N transfer was not only associated with high mycelium density in soil but also with low soil microbial carbon, suggesting that the effect of mycorrhizal fungi on soil microbial populations may be an important factor affecting N transfer between mycorrhizal plants.     

Plant and Soil N Response of Southern Californian Semi-arid Shrublands After 1 Year of Experimental N Deposition

Large inputs of atmospheric N from dry deposition accumulate on vegetation and soil surfaces of southern Californian chaparral and coastal sage scrub (CSS) ecosystems during the late-summer and early-fall and become available as a pulse following winter rainfall; however, the fate of this dry season atmospheric N addition is unknown. To assess the potential for dry season atmospheric N inputs to be incorporated into soil and/or vegetation N pools, an in situ N addition experiment was initiated in a post-fire chaparral and a mature CSS stand where 10 × 10 m plots were exposed to either ambient N deposition (control) or ambient +50 kg N ha⁻¹ (added N) added as NH₄NO₃ during a single application in October 2003. After 1 year of N addition, plots exposed to added N had significantly higher accumulation of extractable inorganic N (NH₄−N + NO₃−N) on ion exchange resins deployed in the 0–10 cm mineral soil layer and higher soil extractable N in the subsurface (30–40 cm) mineral soil than plots exposed to ambient N. Chaparral and CSS shrubs exposed to added N also exhibited a significant increase in tissue N concentration and a decline in the tissue C:N ratio, and added N significantly altered the shrub tissue δ ¹⁵N natural abundance. Leaching of inorganic N to 1 m below the soil surface was on average 2–3 times higher in the added N plots, but large within treatment variability cause these differences to be statistically insignificant. Although a large fraction of the added N could not be accounted for in the shrub and soil N pools investigated, these observations suggest that dry season N inputs can significantly and rapidly alter N availability and shrub tissue chemistry in Mediterranean-type chaparral and CSS shrublands of southern California.     

Litter N retention over winter for a low and a high phenolic species in the alpine tundra

Mycorrhizal fungus colonization of roots may modify plant metal acquisition and tolerance. In the present study, the contribution of the extraradical mycelium of an arbuscular mycorrhizal (AM) fungus, Glomus mosseae (BEG 107), to the uptake of metal cations (Cu, Zn, Cd and Ni) by cucumber (Cucumis sativus) plants was determined. The influence of the amount of P supplied to the hyphae on the acquisition and partitioning of metal cations in the mycorrhizal plants was also investigated. Pots with three compartments were used to separate root and root-free hyphal growing zones. The shoot concentration of Cd and Ni was decreased in mycorrhizal plants compared to non-mycorrhizal plants. In contrast, shoot Zn and Cu concentrations were increased in mycorrhizal plants. High P supply to hyphae resulted in decreased root Cu concentrations and shoot Cd and Ni concentrations in mycorrhizal plants. These results confirm that some elements required for plant growth (P, Zn, Cu) are taken up by mycorrhizal hyphae and are then transported to the plants. Conversely, Cd and Ni were transported in much smaller amounts by hyphae to the plant, so that arbuscular mycorrhizal fungus colonization could partly protect plants from toxic effects of these elements. Selective uptake and transport of plant essential elements over non-essential elements by AM hyphae, increased growth of mycorrhizal plants, and metal accumulation in the root may all contribute to the successful growth of mycorrhizal plants on metal-rich substrates. These effects are stimulated when hyphae can access sufficient P in soil.     

Contribution of an arbuscular mycorrhizal fungus to the uptake of cadmium and nickel in bean and maize plants

Two experiments were carried out in pots with three compartments, a central one for root and hyphal growth and two outer ones which were accessible only for hyphae of the arbuscular mycorrhizal fungus, Glomus mosseae ([Nicol. and Gerd.] Gerdemann and Trappe). In the first experiment, mycorrhizal and nonmycorrhizal bean (Phaseolus vulgaris L.) plants were grown in two soils with high geogenic cadmium (Cd) or nickel (Ni) contents. In the second experiment, mycorrhizal and nonmycorrhizal maize (Zea mays L.) or bean plants were grown in a non-contaminated soil in the central compartment, and either the Cd- or Ni-rich soil in the outer compartments. In additional pots, mycorrhizal plants were grown without hyphal access to the outer compartments. Root and shoot dry weight was not influenced by mycorrhizal inoculation, but plant uptake of metals was significantly different between mycorrhizal and nonmycorrhizal plants. In the first experiment, the contribution of mycorrhizal fungi to plant uptake accounted for up to 37% of the total Cd uptake by bean plants, for up to 33% of the total copper (Cu) uptake and up to 44% of the total zinc (Zn) uptake. In contrast, Ni uptake in shoots and roots was not increased by mycorrhizal inoculation. In the second experiment, up to 24% of the total Cd uptake and also up to 24% of the total Cu uptake by bean could be attributed to mycorrhizal colonisation and delivery by hyphae from the outer compartments. In maize, the mycorrhizal colonisation and delivery by hyphae accounted for up to 41% of the total Cd uptake and 19% of the total Cu uptake. Again, mycorrhizal colonisation did not contribute to Ni uptake by bean or maize. The results demonstrate that the arbuscular mycorrhizal fungus contributed substantially not only to Cu and Zn uptake, but also to uptake of Cd (but not Ni) by plants from soils rich in these metal cations. Deceased 21 September 1996 Deceased 21 September 1996     

Increased drought tolerance of mycorrhizal onion plants caused by improved phosphorus nutrition

Onion plants (Allium cepa L, cv. Downing Yellow Globe) grown in pots and infected by the mycorrhizal fungusGlomus etunicatus Becker and Gerdemann were more drought tolerant than were non-mycorrhizal individials when exposed to several periods of soil water stress separated by periods of high water supply, as shown by greater fresh and dry weights and higher tissue phosphorus levels in the mycorrhizal plants. The tissues of stressed, non-mycorrhizal plants were deficient in P, despite the fact that only non-mycorrhizal plants were fertilized with high levels of P (26 mg P per 440 g soil). Differences in plant water relations (leaf water potentials or transpiration rates) and changes in soil P levels which may have affected plant growth were investigated, and discounted as factors important for the results. The P nutrition of plants has been implicated in the ability of plants to tolerate drought and it was concluded that the ability of the mycorrhizal fungus to maintain adequate P nutrition in the onions during soil water stress was a major factor in the improved drought tolerance. Infection of the root by the fungus was found not to be affected by water stress or P fertilization but fungal reproduction, as determined by spore numbers in the soil, was decreased by water stress and by P fertilization.Michigan Agricultural Experiment Station Article No. 10050     

The effect of drought on mycorrhizas of beech (Fagus sylvatica L.): changes in community structure,and the content of carbohydrates and nitrogen storage bodies of the fungi

In a water-exclusion experiment, five different ecotypes of beech (Fagus sylvatica L.; representing regions of different environmental and climatic conditions in Baden-Württemberg, Germany) were subjected to drought conditions of different severity between July and September of two consecutive years. Drought stress as characterised by the water content and the pre-dawn water potential of the leaves was related to the degree of mycorrhization, the type of ectomycorrhiza, and the physiological properties of individual fungus/plant interactions at the fine roots of different beech ecotypes. Our data show that decreased soil water availability did not significantly change either the degree of fungal colonisation of beech roots (measured by the amount of ergosterol) or the number of ectomycorrhizal types per root system. Drought did, however, have an influence on the composition of the ectomycorrhizal community, and different mycorrhizal types responded to drought differently in terms of their patterns of occurrence/abundance. While the abundance of the dominant mycorrhizal types, formed with Byssocorticium atrovirens and Lactarius subdulcis, was not affected, drought increased the abundance of mycorrhiza formed between beech and Xerocomus chrysenteron. A detailed analysis of plant and fungal carbohydrates in mycorrhizas indicated that different drought intensities led to distinguishable responses. In plants exhibiting a pre-dawn water potential of down to -1.96 MPa, drought caused the accumulation of sucrose, glucose and fructose, and of fungus-specific compounds such as mannitol and arabitol in mycorrhizal roots at the expense of, e.g. trehalose. The accumulation of sugar alcohols, which constitute compatible solutes known to counteract drought stress, was species-specific. Mycorrhizas with X. chrysenteron formed large amounts of arabitol, while those with L. subdulcis accumulated mannitol. Sustained partitioning of carbon towards the mycorrhizal fungi under drought was also reflected by an increase of nitrogen storage in the fungal vacuoles. In treatments where the pre-dawn water potential reached values of as low as -2.4 MPa, such alterations were no longer found. In such plants, the starch and soluble sugars content was generally reduced, which also resulted in a lack of increase in protective, fungus-specific sugar alcohols. In summary, the data show that, within certain limits, an increase in drought causes a shift in plant/fungus communities. The shift in the pattern of fungus-specific compounds could possibly be used as a sensitive measure of physiological stress imposed on this symbiosis.     

The proliferation of fungal hyphae in soils supporting mycorrhizal and non-mycorrhizal plants

This study investigated the impact of mycorrhizal plants, non-mycorrhizal plants and soil organic matter on the relative abundance of soil hyphae perceived to belong to indigenous arbuscular mycorrhizal (AM) plants. The mycorrhizal plants corn (Zea mays L.) and barley (Hordeum vulgare L.) and a non-mycorrhizal plant, canola (Brassica napus L.), were grown in unsterilized soil in pots inoculated with mycorrhizal corn root fragments. The abundance of hyphae was measured after 5 weeks and the response of fungal growth to the addition of corn residues in the absence of plants was assessed. The abundance of hyphae was higher in the presence of the mycorrhizal plants than in the other treatments. AM hyphae present under mycorrhizal plants accounted for more than 83% of the measured hyphae. The levels of root colonization of 32% in corn and 27% in barley confirmed the mycorrhizal status of the experimental plants. Only a few points of entry were observed in canola, the non-host plant. The percentage of mycorrhizal colonization was positively related (R ²?=?0.85) to the abundance of soil hyphae, indicating that AM hyphae were the major component of the soil hyphae in the presence of mycorrhizal plants in this study.     

Response ofBromus inermis inoculated withGlomus fasciculatum to potassium fertilization and drought stress

Summary Bromus inermis Leyss. was grown in a 2×2×2 factorial design using different levels of mycorrhizal inoculation (inoculated and noninoculated), soil water stress (Ψ₁ or −0.8 MPa) and potassium (K) fertilization (0 or 150 ppm) as factors. Soil water stress and mycorrhizal inoculation significantly reduced plant top dry weight during the 18 week study. Chlamydospore production by the mycorrhizal symbiontGlomus fasciculatum (Thaxter sensu. Gerd.) Gerd. and Trappe was significantly reduced by soil water stress of −0.8 MPa. Potassium (K) fertilization did not significantly influence plant top growth or mycorrhizal colonization. However, foliar Ca and Mg were significantly lower in plants fertilized with K. Foliar Ca and Mg concentrations of P, K, N, Mn, Zn and Cu were significantly greater in drought stressed plants whereas Ca and Mg concentrations were significantly greater in well-watered plants.     

Uptake and transport of organic and inorganic nitrogen by arbuscular mycorrhizal fungi

New information on N uptake and transport of inorganic and organic N in arbuscular mycorrhizal fungi is reviewed here. Hyphae of the arbuscular mycorrhizal fungus Glomus mosseae (Nicol. and Gerd.) Gerd. and Trappe (BEG 107) were shown to transport N supplied as ¹⁵N-Gly to wheat plants after a 48 h labelling period in semi-hydroponic (Perlite), non-sterile, compartmentalised pot cultures. Of the ¹⁵N supplied to hyphae in pot cultures over 48 h, 0.2 and 6% was transported to plants supplied with insufficient N or sufficient N, respectively. The increased ¹⁵N uptake at the higher N supply was related to the higher hyphal length density at the higher N supply. These findings were supported by results from in vitro and monoxenic studies. Excised hyphae from four Glomus isolates (BEG 84, 107, 108 and 110) acquired N from both inorganic (¹⁵NH₄ ¹⁵NO₃, ¹⁵NO₃ ⁻ or ¹⁵NH₄ ⁺) and organic (¹⁵N-Gly and ¹⁵N-Glu, except in BEG 84 where amino acid uptake was not tested) sources in vitro during short-term experiments. Confirming these studies under sterile conditions where no bacterial mineralisation of organic N occurred, monoxenic cultures of Glomus intraradices Schenk and Smith were shown to transport N from organic sources (¹⁵N-Gly and ¹⁵N-Glu) to Ri T-DNA transformed, AM-colonised carrot roots in a long-term experiment. The higher N uptake (also from organic N) by isolates from nutrient poor sites (BEG 108 and 110) compared to that from a conventional agricultural field implied that ecotypic differences occur. Although the arbuscular mycorrhizal isolates used contributed to the acquisition of N from both inorganic and organic sources by the host plants/roots used, this was not enough to increase the N nutritional status of the mycorrhizal compared to non-mycorrhizal hosts. This revised version was published online in June 2006 with corrections to the Cover Date.     

Hyphal N transport by a vesicular-arbuscular mycorrhizal fungus associated with cucumber grown at three nitrogen levels

Cucumis sativus L. cv. Aminex (F1 hybrid) was grown alone or in symbiosis with Glomus intraradices Schenck and Smith in containers with two hyphal compartments (HC_A and HC_B) on either side of a root compartment (RC) separated by fine nylon mesh. Plants received a total of either 100, 200 or 400 mg N which were applied gradually to the RC during the experiment. ¹⁵N was supplied to HC_A 42 d after plating, at 50 mg ¹⁵NH₄ ⁺-N kg^–1 soil. Lateral movement of the applied ¹⁵N towards the roots was minimized by using a nitrification inhibitor and a hyphal buffer compartment.Non-mycorrhizal controls contained only traces of ¹⁵N after a 27 d labelling period irrespective of the amount of N supplied to the RC. In contrast, 49, 48 and 27% of the applied ¹⁵N was recovered in mycorrhizal plants supplied with 100, 200 and 400 mg N, respectively. The plant dry weight was increased by mycorrhizal colonization at all three levels of N supply, but this effect was strongest in plants of low N status. The results indicated that this increase was due partly to the improved inflow of N via the external hyphae. Root colonization by G. intraradices was unaffected by the amount of N supplied to the RC, while hyphal length increased in HC_A compared to HC_B. Although a considerable ¹⁵N content was detected in mycorrhizal roots adjacent to HC_B, only insignificant amounts of ¹⁵N were found in the external hyphae in HC_B. The external hyphae depleted the soil of inorganic N in both HC_A and HC_B, while the concentration of soil mineral N was still high in non-mycorrhizal containers at harvest. An exception was plants supplied with 400 mg N, where some inorganic N was present at 5 cm distance from the RC in HC_A. The possibility of a regulation mechanism for hyphal transport of N is discussed.     

Seasonal response to drought and rewatering in Portulacaria afra (L.) Jacq.

Summary Gas exchange characteristics of droughted and rewatered Portulacaria afra were studied during the seasonal shift from CAM to C₃ photosynthesis. ¹⁴CO₂ uptake, stomatal conductance, and total titratable acidity were determined for both irrigated and 2, 4, and 7.5 month waterstressed plants from summer 1984 to summer 1985. Irrigated P. afra plants were utilizing the CAM pathway throughout the summer and shifted to C₃ during the winter and spring. Beginning in September, P. afra plants shifted from CAM to CAM-idling after 2 months of water-stress. When water-stress was initiated later in the fall, exogenous CO₂ uptake was still measurable after 4 months of drought. After 7.5 months of stress, exogenous CO₂ uptake was absent. The shift from CAM to CAM-idling or C₃ in the fall and winter was related to when water stress was initiated and not to the duration of the stress. Gas exchange resumed within 24 h of rewatering regardless of the duration of the drought. In the winter and spring, rewatering resulted in a full resumption of daytime CO₂ uptake. Whereas during the summer, rewatering quickly resulted in early morning CO₂ uptake, but nocturnal CO₂ uptake through the CAM pathway was observed after 7 days. Gas exchange measurements, rewatering characteristics, and transpirational water loss support the hypothesis that the C₃ pathway was favored during the winter and spring. The CAM pathway was functional during the summer when potential for water loss was greater. Our investigations indicate that P. afra has a flexible photosynthetic system that can withstand long-term drought and has a rapid response to rewatering.     

N-P fertilization did not reduce AMF abundance or diversity but alter AMF composition in an alpine grassland infested by a root hemiparasitic plant

Fertilization has been shown to have suppressive effects on arbuscular mycorrhizal fungi (AMF) and root hemiparasites separately in numerous investigations, but its effects on AMF in the presence of root hemiparasites remain untested. In view of the contrasting nutritional effects of AMF and root hemiparasites on host plants, we tested the hypothesis that fertilization may not show strong suppressive effects on AMF when a plant community was infested by abundant hemiparasitic plants. Plants and soil samples were collected from experimental field plots in Bayanbulak Grassland, where N and P fertilizers had been applied for three continuous years for control against a spreading root hemiparasite, Pedicularis kansuensis. Shoot and root biomass of each plant functional group were determined. Root AMF colonization levels, soil spore abundance, and extraradical hyphae length density were measured for three soil depths (0-10 cm, 10-20 cm, 20-30 cm). Partial 18S rRNA gene sequencing was used to detect AMF diversity and community composition. In addition, we analyzed the relationship between relative abundance of different AMF genera and environmental factors using Spearman's correlation method. In contrast to suppressive effects reported by many previous studies, fertilization showed no significant effects on AMF root colonization or AMF species diversity in the soil. Instead, a marked increase in soil spore abundance and extraradical hyphae length density were observed. However, fertilization altered relative abundance and AMF composition in the soil. Our results support the hypothesis that fertilization does not significantly influence the abundance and diversity of AMF in a plant community infested by P. kansuensis.     

Acquisition of phosphorus and copper by VA-mycorrhizal hyphae and root-to-shoot transport in white clover

White clover (Trifolium repens L.) plants were grown in a calcareous soil in pots with three compartments, a central one for root growth and two outer ones for growth of vesicular-arbuscular (VA) mycorrhizal (Glomus mosseae [Nicol. & Gerd.] Gerdemann & Trappe) hyphae (hyphal compartments). Phosphorus (P) was applied at three levels (0, 20 and 50 mg kg⁻¹ soil) in the outer compartments in mycorrhizal treatments. Root and shoot dry weight were increased in mycorrhizal plants with hyphal access to outer compartments. Growth of the mycorrhizal hyphae in the outer compartments was not significantly affected by variation in P level in these compartments. However, both concentration and amount of P in roots and shoots sharply increased with increasing P supply in the outer (hyphal) compartments. With increasing P levels the calculated delivery of P by the hyphae from the outer compartments increased from 34% to 90% of total P uptake. Hyphal access to the outer compartments also significantly increased both concentration and quantity of Cu in the plants. The calculated delivery of Cu by the hyphae from the outer compartments ranged from 53% to 62% of total Cu uptake, irrespective of the P levels and the amounts of P taken up and transported by the hyphae. However, the distribution of Cu over roots and shoots was largely dependent on P levels. With increase in P level in the outer compartments the calculated hyphal contribution to the total amount of Cu in the shoots increased from 12% to 58%, but decreased in the roots from 75% to 46%. In conclusion, uptake and transport by VA-mycorrhizal hyphae may contribute substantially not only to P nutrition, but also to Cu nutrition of the host.     

WATER RELATIONS OF TWO CALIFORNIA CHAPARRAL SHRUBS

Water use patterns of two California chaparral shrub species, chamise (Adenostoma fasciculatum H. and A.) and Stanford manzanita (Arctostaphylos stanfordiana Parry), were compared during summer drought. Observations of diurnal and seasonal courses of shoot water potential, leaf conductance and transpiration revealed that chamise was more conservative in water use than manzanita. Evidence obtained cast doubt on a hypothesis previously proposed to explain an anomalous pattern of shoot water potential in chamise.