Mycorrhizal fungi enhance plant nutrient acquisition and modulate nitrogen loss with variable water regimes

Climate change will alter both the amount and pattern of precipitation and soil water availability, which will directly affect plant growth and nutrient acquisition, and potentially, ecosystem functions like nutrient cycling and losses as well. Given their role in facilitating plant nutrient acquisition and water stress resistance, arbuscular mycorrhizal (AM) fungi may modulate the effects of changing water availability on plants and ecosystem functions. The well‐characterized mycorrhizal tomato (Solanum lycopersicum L.) genotype 76R (referred to as MYC+) and the mutant mycorrhiza‐defective tomato genotype rmc were grown in microcosms in a glasshouse experiment manipulating both the pattern and amount of water supply in unsterilized field soil. Following 4 weeks of differing water regimes, we tested how AM fungi affected plant productivity and nutrient acquisition, short‐term interception of a ${}^{15}{\text{NH}}_4^{+}$ pulse, and inorganic nitrogen (N) leaching from microcosms. AM fungi enhanced plant nutrient acquisition with both lower and more variable water availability, for instance increasing plant P uptake more with a pulsed water supply compared to a regular supply and increasing shoot N concentration more when lower water amounts were applied. Although uptake of the short‐term ${}^{15}{\text{NH}}_4^{+}$ pulse was higher in rmc plants, possibly due to higher N demand, AM fungi subtly modulated ${\text{NO}}_3^{?}$ leaching, decreasing losses by 54% at low and high water levels in the regular water regime, with small absolute amounts of ${\text{NO}}_3^{?}$ leached (<1 kg N/ha). Since this study shows that AM fungi will likely be an important moderator of plant and ecosystem responses to adverse effects of more variable precipitation, management strategies that bolster AM fungal communities may in turn create systems that are more resilient to these changes.     

Fertilization alters nitrogen isotopes and concentrations in ectomycorrhizal fungi and soil in pine forests

To assess how nitrogen (N) availability affected ectomycorrhizal functioning and to test a theoretical model of ectomycorrhizal ¹⁵N partitioning, we measured C/N and δ¹⁵N in soils and nine fungal taxa in two Swedish N addition experiments. Sporocarp C/N and soil C/N decreased with fertilization, implying that N uptake per unit fungal growth increased. The S horizon was more responsive than the F and H horizons to changes in N addition, with N turnover for these horizons of 24, 57, and 57 y, respectively. Fungal and soil δ¹⁵N patterns identified fungal N sources, with N acquisition primarily from the S, F, or H horizon for two, five, and two taxa, respectively. With increasing N availability, sporocarp ¹⁵N enrichment increased in five taxa, in agreement with our model of fungal-plant N partitioning. However, it decreased in Lactarius rufus and Russula aeruginea, perhaps indicating shifts towards greater inorganic N uptake in these two taxa. This may relate to the generally lower sensitivity of these taxa to N deposition compared to the Cortinarius and Suillus taxa that fit our model of ¹⁵N partitioning.     

丛枝菌根真菌与氮添加对不同根形态基因型水稻氮吸收的影响

植物主要依赖自身根系从土壤中获取矿质养分; 具有不同根形态的植物对于养分的吸收能力存在差异。丛枝菌根真菌(AMF)能与陆地植物根系形成共生关系, 帮助植物吸收矿质养分。但是, AMF对于植物根系养分吸收的促进效应是否会受根形态的影响还鲜有研究。该研究选取4种不同根形态基因型水稻(根毛缺陷突变体rhl1、侧根缺陷突变体iaa11、不定根缺失突变体arl1和野生型Kas)为研究对象, 设置2种施氮水平处理(低氮: 20 mg·kg^-1氨氮; 高氮: 100 mg·kg^-1氨氮), 利用稳定同位素¹⁵N示踪标记技术, 探究AMF和氮添加对不同根形态植物氮吸收的影响。研究结果发现, 相比低氮处理, 高氮处理下, rhl1、Kas、iaa11与arl1的茎叶¹⁵N浓度分别提高了60%、72%、128%与118%, 说明氮添加显著促进了水稻氮吸收, 且iaa11与arl1对氮添加的响应更强烈。在低氮水平下, AMF对rhl1、Kas、iaa11与arl1氮吸收的平均效应值分别为17%、31%、42%、51%, 表明AMF对于植物氮吸收的促进效应受根形态影响, iaa11与arl1对AMF的响应明显高于Kas与rhl1; 相较于低氮水平, 高氮水平下AMF对于不同根形态水稻氮吸收的促进效应都会显著降低, 表明氮添加削弱了AMF对植物氮吸收的促进效应。该研究阐明了4种不同根形态基因型水稻氮养分吸收存在显著差异, 其中氮吸收能力较弱的基因型水稻对AMF的响应更强, 该结果补充了植物与AMF在养分吸收上存在功能互补的控制实验证据。     

Effects of simultaneous infections of endophytic fungi and arbuscular mycorrhizal fungi on the growth of their shared host grass Achnatherum sibiricum under varying N and P supply

Grasses can be infected by endophytic fungi and arbuscular mycorrhizal (AM) fungi simultaneously. Here, we investigated the interactions of a native grass, Achnatherum sibiricum, with both endophytic and AM fungi (Glomus mosseae, GM and Glomus etunicatum, GE) at different nitrogen (N) and phosphorus (P) levels. The results showed that endophyte infection significantly suppressed the colonization rates and spore density of GE, but had no effect on those of GM. Endophyte infection increased shoot biomass regardless of the nutrient conditions. The effects of AM fungi on host growth were dependent on mycorrhizal species. There was no significant interaction between endophytic fungi and GE on host growth; however, a significant interaction between endophytic fungi and GE occurred in total phenolic concentrations and P uptake. As for GM, a significant interaction among endophytic fungi, AM fungi and nutrient availability occurred in shoot growth. Under sufficient N and P conditions, endophyte infection alleviated the detrimental effects of GM colonization on host growth.     

施肥对垂穗披碱草根系中丛枝菌根真菌的影响

采用传统染色与克隆测序的方法,研究了8年不同施肥(氮磷)梯度对垂穗披碱草根系中丛枝菌根(AM)侵染率和AM真菌群落的影响.结果表明: 随施肥浓度升高, 垂穗披碱草根系单位根长AM总侵染率从67.5%下降至7.3%,丛枝侵染率从5.2%降至0.1%.根系共检测出24个AM真菌分子种,但随着施肥浓度上升,AM真菌的平均物种丰富度从6种下降至2.6种.不同施肥处理对AM真菌群落结构有显著影响,土壤速效磷和根系氮含量与AM真菌群落呈极显著相关.氮磷有效性随施肥梯度逐渐上升,且与AM侵染率和AM真菌物种丰富度呈显著负相关.施高浓度氮磷肥对AM共生体有明显的抑制作用,导致AM真菌物种多样性丧失.     

Arbuscular mycorrhizal fungi can transfer substantial amounts of nitrogen to their host plant from organic material

Nitrogen (N) capture by arbuscular mycorrhizal (AM) fungi from organic material is a recently discovered phenomenon. This study investigated the ability of two Glomus species to transfer N from organic material to host plants and examined whether the ability to capture N is related to fungal hyphal growth. Experimental microcosms had two compartments; these contained either a single plant of Plantago lanceolata inoculated with Glomus hoi or Glomus intraradices, or a patch of dried shoot material labelled with (15)N and (13)carbon (C). In one treatment, hyphae, but not roots, were allowed access to the patch; in the other treatment, access by both hyphae and roots was prevented. When allowed, fungi proliferated in the patch and captured N but not C, although G. intraradices transferred more N than G. hoi to the plant. Plants colonized with G. intraradices had a higher concentration of N than controls. Up to one-third of the patch N was captured by the AM fungi and transferred to the plant, while c. 20% of plant N may have been patch derived. These findings indicate that uptake from organic N could be important in AM symbiosis for both plant and fungal partners and that some AM fungi may acquire inorganic N from organic sources.     

The influence of Rhizobium and arbuscular mycorrhizal fungi on nitrogen and phosphorus accumulation by Vicia faba

BACKGROUND AND AIMS: The aim of this study was to investigate the effects of the interactions between the microbial symbionts, Rhizobium and arbuscular mycorrhizal fungi (AMF) on N and P accumulation by broad bean (Vicia faba) and how increased N and P content influence biomass production, leaf area and net photosynthetic rate. METHODS: A multi-factorial experiment consisting of four different legume-microbial symbiotic associations and two nitrogen treatments was used to investigate the influence of the different microbial symbiotic associations on P accumulation, total N accumulation, biomass, leaf area and net photosynthesis in broad bean grown under low P conditions. KEY RESULTS: AMF promoted biomass production and photosynthetic rates by increasing the ratio of P to N accumulation. An increase in P was consistently associated with an increase in N accumulation and N productivity, expressed in terms of biomass and leaf area. Photosynthetic N use efficiency, irrespective of the inorganic source of N (e.g. NO3- or N2), was enhanced by increased P supply due to AMF. The presence of Rhizobium resulted in a significant decline in AMF colonization levels irrespective of N supply. Without Rhizobium, AMF colonization levels were higher in low N treatments. Presence or absence of AMF did not have a significant effect on nodule mass but high N with or without AMF led to a significant decline in nodule biomass. Plants with the Rhizobium and AMF symbiotic associations had higher photosynthetic rates per unit leaf area. CONCLUSIONS: The results indicated that the synergistic or additive interactions among the components of the tripartite symbiotic association (Rhizobium-AMF-broad bean) increased plant productivity.     

Carbon for nutrient exchange between arbuscular mycorrhizal fungi and wheat varies according to cultivar and changes in atmospheric carbon dioxide concentration

Arbuscular mycorrhizal fungi (AMF) form symbioses with most crops, potentially improving their nutrient assimilation and growth. The effects of cultivar and atmospheric CO₂ concentration ([CO₂]) on wheat–AMF carbon‐for‐nutrient exchange remain critical knowledge gaps in the exploitation of AMF for future sustainable agricultural practices within the context of global climate change. We used stable and radioisotope tracers (¹⁵N, ³³P, ¹⁴C) to quantify AMF‐mediated nutrient uptake and fungal acquisition of plant carbon in three wheat (Triticum aestivum L.) cultivars. We grew plants under current ambient (440 ppm) and projected future atmospheric CO₂ concentrations (800 ppm). We found significant ¹⁵N transfer from fungus to plant in all cultivars, and cultivar‐specific differences in total N content. There was a trend for reduced N uptake under elevated atmospheric [CO₂]. Similarly, ³³P uptake via AMF was affected by cultivar and atmospheric [CO₂]. Total P uptake varied significantly among wheat cultivars and was greater at the future than current atmospheric [CO₂]. We found limited evidence of cultivar or atmospheric [CO₂] effects on plant‐fixed carbon transfer to the mycorrhizal fungi. Our results suggest that AMF will continue to provide a route for nutrient uptake by wheat in the future, despite predicted rises in atmospheric [CO₂]. Consideration should therefore be paid to cultivar‐specific AMF receptivity and function in the development of climate smart germplasm for the future.     

Symbiotic soil fungi enhance ecosystem resilience to climate change

Substantial amounts of nutrients are lost from soils through leaching. These losses can be environmentally damaging, causing groundwater eutrophication and also comprise an economic burden in terms of lost agricultural production. More intense precipitation events caused by climate change will likely aggravate this problem. So far it is unresolved to which extent soil biota can make ecosystems more resilient to climate change and reduce nutrient leaching losses when rainfall intensity increases. In this study, we focused on arbuscular mycorrhizal (AM) fungi, common soil fungi that form symbiotic associations with most land plants and which increase plant nutrient uptake. We hypothesized that AM fungi mitigate nutrient losses following intensive precipitation events (higher amount of precipitation and rain events frequency). To test this, we manipulated the presence of AM fungi in model grassland communities subjected to two rainfall scenarios: moderate and high rainfall intensity. The total amount of nutrients lost through leaching increased substantially with higher rainfall intensity. The presence of AM fungi reduced phosphorus losses by 50% under both rainfall scenarios and nitrogen losses by 40% under high rainfall intensity. Thus, the presence of AM fungi enhanced the nutrient interception ability of soils, and AM fungi reduced the nutrient leaching risk when rainfall intensity increases. These findings are especially relevant in areas with high rainfall intensity (e.g., such as the tropics) and for ecosystems that will experience increased rainfall due to climate change. Overall, this work demonstrates that soil biota such as AM fungi can enhance ecosystem resilience and reduce the negative impact of increased precipitation on nutrient losses.     

丛枝菌根真菌和根瘤菌对白三叶氮同化的影响

为揭示丛枝菌根真菌(AMF)和根瘤菌在白三叶氮(N)同化中的作用,该研究对白三叶进行单一或联合接种隐类球囊霉(Paraglomus occultum)和三叶草根瘤菌(Rhizobium trifolii),分析其对白三叶的生长、光合作用、叶片N和氨基酸含量以及N同化相关酶活性的影响。结果表明:(1)单一接种AMF或根瘤菌以及联合接种AMF和根瘤菌均显著增加了白三叶的株高、匍匐茎长度、叶片数、地上部生物量、总生物量、叶绿素b和总叶绿素含量、稳态光量子效率和叶片N含量,这种增强效应是联合接种>单一AMF>单一根瘤菌>未接种处理。(2)联合接种AMF和根瘤菌显著增加了白三叶叶片中丙氨酸、精氨酸、天冬酰胺、天冬氨酸、谷氨酰胺、谷氨酸和组氨酸的含量,显著提升了叶片N同化相关酶如硝酸还原酶、亚硝酸还原酶、谷氨酰胺合成酶、谷氨酸合成酶、谷氨酸脱氢酶、天冬酰胺合成酶和天冬氨酸转氨酶的活性,显著促进AMF对白三叶根系的侵染。综上认为,联合接种AMF和根瘤菌通过激活N同化相关酶活性有效促进N同化,产生更多氨基酸,进一步促进白三叶植株生长; 联合接种AMF和根瘤菌具有协同作用,有效促进了白三叶的N同化。     

丛枝菌根真菌在内的土壤真菌对入侵植物紫茎泽兰的正反馈：证据来自野外实验

包括紫茎泽兰在内的许多外来植物都能够与新入侵生境的丛枝菌根真菌（ AMF）形成互利共生,因此菌根真菌如何调节外来植物种的入侵是当前亟待研究的问题。测定了紫茎泽兰入侵不同阶段（紫茎泽兰呈零星丛状分布于本地植物群落中[部分入侵生境]及紫茎泽兰单优群落形成期[入侵生境]）的土壤化学性状,而后通过野外试验,采用杀真菌剂处理,研究了包括AMF在内的土壤真菌对紫茎泽兰入侵的反馈作用。紫茎泽兰入侵改变了土壤化学性状。施用杀真菌剂降低了紫茎泽兰叶面积、叶片碳、氮、磷、和δ13 C含量。综合分析发现,在紫茎泽兰与本地植物混生群落中,土壤真菌能够增加紫茎泽兰叶片碳和δ13 C含量,但是不能提高紫茎泽兰的光合作用,表明碳和δ13 C含量的提高,不是光合作用的结果,而是通过其他机制实现的。因此可以得出,在部分入侵生境中,碳从土壤或临近植物经由菌丝网向紫茎泽兰转移。紫茎泽兰入侵不同阶段土壤养分的变化利于紫茎泽兰种群建立,同时利于紫茎泽兰借助真菌（尤其是AMF）从土壤或临近植物转移碳,促进种群扩散,这可能是紫茎泽兰入侵的机制之一。     

氮和土著AMF对黄瓜间作土壤酶活性及氮利用的影响

设施土壤氮（N）肥的大量不合理施用和高残留是导致作物硝态N含量超标和农业面源污染的主要因素之一。研究土著丛枝菌根真菌（arbuscular mycorrhizal fungi,AMF）与间作体系强化蔬菜对不同形态N的利用并结合土壤酶活性的反馈作用,可为设施土壤N素的高效利用和降低土壤N残留提供依据。本研究采用盆栽试验,设置黄瓜单作和黄瓜//大豆间作种植模式,不同AMF处理[不接种（NM）、接种土著AMF]和不同形态N处理[不施N（N0）、有机N（谷氨酰胺120mg/kg,ON120）、无机N（碳酸氢铵120mg/kg,ION120）],探讨了设施条件下施用不同形态N、接种土著AMF与间作大豆对黄瓜根围土壤酶活性及氮利用的影响。结果表明,与NM相比,接种土著AMF使设施黄瓜地上部、根系生物量及植株N吸收量均有不同程度的增加,根围土壤NH₄ ⁺-N、NO₃ ^--N含量呈现降低趋势。同一N处理-土著AMF条件下,间作大豆处理下的黄瓜根系菌根侵染率显著高于单作处理;间作大豆也使黄瓜植株地上部、根系生物量及N吸收量显著增加,同时显著降低了根围土壤铵态N含量。此外,间作-土著AMF条件下,ON120和ION120处理的黄瓜根围土壤脲酶活性较N0处理分别提高了30%和14%,蛋白酶和硝酸还原酶活性也呈现出相同趋势。可见,所有复合处理中,以间作体系接种土著AMF与施用适量有机N的组合明显促进了设施黄瓜生长和N素利用率。     

Arbuscular mycorrhizal fungi in the tree seedlings of two Australian rain forests: occurrence, colonization, and relationships with plant performance

The roots of rain forest plants are frequently colonized by arbuscular mycorrhizal fungi (AMF) that can promote plant growth in the nutrient poor soils characteristic of these forests. However, recent studies suggest that both the occurrence of AMF on rain forest plants and the dependence of rain forest plants on AMF can be highly variable. We examined the occurrence and levels of AMF colonization of some common seedling species in a tropical and a subtropical rain forest site in Queensland, Australia. We also used a long-term database to compare the growth and mortality rates of seedling species that rarely formed AMF with those that regularly formed AMF. In both forests, more than one-third of the seedling species rarely formed AMF associations, while 40% of species consistently formed AMF in the tropical site compared to 27% in the subtropical site. Consistent patterns of AMF occurrence were observed among plant families at the two sites. Variation among seedling species in AMF occurrence or colonization was not associated with differences in seed mass among species, variation in seedling size and putative age within a species, or lack of AMF inoculum in the soil. Comparisons of four seedling species growing both in the shaded understory and in small canopy gaps revealed an increase in AMF colonization in two of the four species in gaps, suggesting that light limitation partially explains the low occurrence of AMF. Seedling survival was significantly positively associated with seed biomass but not with AMF colonization. Furthermore, seedling species that regularly formed AMF and those that did not had similar rates of growth and survival, suggesting that mycorrhizal and nonmycorrhizal strategies were equivalent in these forests. Furthermore, the high numbers of seedlings that lacked AMF and the overall low rate of seedling growth (the average seedling required 6 years to double its height) suggest that most seedlings did not receive significant indirect benefits from AMF through connection to canopy trees via a common mycorrhizal network.