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1.
土壤盐渍化是世界性的生态环境问题之一。关于丛枝菌根真菌(AM真菌)对盐胁迫下白芨生长和养分吸收的影响研究相对较少。以白芨幼苗为研究对象,设置盐处理(0 mM和200 mM)和AM真菌处理(不接种AM真菌和接种摩西球囊霉)各两个水平共计4个处理组合,探讨盐胁迫下AM真菌对白芨幼苗生长和养分吸收的影响。结果表明,盐胁迫显著降低白芨根系侵染率、总干重和地上部分P含量分别约56.1%、48.9%和10%;在对照盐浓度下,接种AM真菌显著提高了白芨根系侵染率、总干重、地上部分N含量及地上部分P含量分别约14%、56.1%、10.8%和8.2%而在高盐浓度下,接种AM真菌则显著提高白芨根系侵染率、总干重、地上部分N含量及地下部分P含量分别约42.7%、30.7%、12.1%和11%,说明接种AM真菌可降低盐胁迫对白芨的生长和养分吸收的抑制作用。以上结果表明,接种AM真菌有助于提高白芨的耐盐能力,这对应用菌根技术提高白芨在盐化生境的种植和管理具有理论和实践指导意义。  相似文献   

2.
丛枝菌根真菌对番茄信号物质的诱导效应   总被引:1,自引:0,他引:1  
王彬  李辉  徐丽娟  李敏  刘润进 《菌物学报》2010,29(4):561-568
盆栽番茄Lycopersicon esculentum幼苗分别接种丛枝菌根(AM)真菌摩西球囊霉Glomus mosseae、地表球囊霉G.versiforme、根内球囊霉G.intraradices、幼套球囊霉G.etunicatum及珠状巨孢囊霉Gigaspora margarita 35d后,开始测定番茄植株内源信号物质水杨酸(SA)、茉莉酸(JA)、一氧化氮(NO)和过氧化氢(H2O2)含量变化,抗性相关酶活性,丙二醛(MDA)含量以及生长量等指标。结果表明,接种AM真菌增加了番茄植株鲜重、株高、地上部和地下部干重、叶片和根系NO、JA、H2O2含量和结合态SA含量,其中,以摩西球囊霉G.mosseae诱导作用最大,叶片和根系内NO、JA、H2O2和结合态SA含量分别比对照增加了3.3和1.9倍、6.8和8.0倍、0.9和1.2倍、1.9和2.6倍,而根系中游离态SA含量一直处于较低水平,只有摩西球囊霉G.mosseae处理在诱导高峰时根系游离态SA含量比对照略有增加。接种AM真菌处理的番茄叶片和根系超氧化物歧化酶(SOD)、过氧化物酶(POD)、过氧化氢酶(CAT)、苯丙氨酸解氨酶(PAL)活性显著增加,其中以摩西球囊霉G.mosseae的诱导效应最大,与未接种对照相比分别增加了0.6和0.3倍、7.9和3.1倍、0.4和1.2倍、2.3和1.9倍;幼套球囊霉G.etunicatum的诱导效应最小:与未接种对照相比分别增加了0.26和0.14倍、2.3和1.0倍、0.1和0.28倍、0.55和0.31倍;而MDA含量下降,分别降低了66%和68%、34%和41%、51%和50%、12%和26%、18%和29%。表明AM真菌能诱导植物同时产生多种信号物质,而且这些信号参与了AM真菌-番茄共生体系统抗性的表达。  相似文献   

3.
水分胁迫下AM真菌对沙打旺生长和抗旱性的影响   总被引:7,自引:0,他引:7  
郭辉娟  贺学礼 《生态学报》2010,30(21):5933-5940
利用盆栽试验研究了水分胁迫条件下接种AM真菌对优良牧草和固沙植物沙打旺(Astragalus adsurgens Pall.)生长和抗旱性的影响。在土壤相对含水量为70%、50%和30%条件下,分别接种摩西球囊霉(Glomus mosseae)和沙打旺根际土著菌,不接种处理作为对照。结果表明,水分胁迫显著降低了沙打旺植株(无论接种AM真菌与否)的株高、分枝数、地上部干重和地下部干重,并显著提高了土著AM真菌的侵染率,对摩西球囊霉的侵染率无显著影响。接种AM真菌可以促进沙打旺生长和提高植株抗旱性,但促进效应因土壤含水量和菌种不同而存在差异。不同水分条件下,接种AM真菌显著提高了植株菌根侵染率、根系活力、地下部全N含量和叶片CAT活性。土壤相对含水量为30%和50%时,接种株地上部全N、叶片叶绿素、可溶性蛋白、脯氨酸含量和POD活性显著高于未接种株;接种AM真菌显著降低了叶片MDA含量;接种土著AM真菌的植株株高、分枝数、地上部和地下部干重显著高于未接种株。土壤相对含水量为30%时,接种AM真菌显著增加了地上部全P含量和叶片相对含水量;接种摩西球囊霉的植株株高、分枝数、地上部和地下部干重显著高于未接种株。水分胁迫40d,接种AM真菌显著提高了叶片可溶性糖含量。水分胁迫80d,接种株叶片SOD活性显著增加。菌根依赖性随水分胁迫程度增加而提高。沙打旺根际土著菌接种效果优于摩西球囊霉。水分胁迫和AM真菌的交互作用对分枝数、菌根侵染率、叶片SOD、CAT和POD活性、叶绿素、脯氨酸、可溶性蛋白、地上部全N和全P、地下部全N和根系活力有极显著影响,对叶片丙二醛和地下部全P有显著影响。AM真菌促进根系对土壤水分和矿质营养的吸收,改善植物生理代谢活动,从而提高沙打旺抗旱性,促进其生长。试验结果为筛选优良抗旱菌种,充分利用AM真菌资源促进荒漠植物生长和植被恢复提供了依据。  相似文献   

4.
为初步探究丛枝菌根(arbuscularmycorrhizal,AM)真菌促进百合生长并延长瓶插过程中切花观赏期的作用机制,于温室盆栽条件下对百合Lilium brownii进行摩西斗管囊霉Funneliformis mosseae和变形球囊霉Glomusversiforme单一接种或者共同双接种处理。结果表明,共同接种F.mosseae和G.versiforme的百合地上部干重和地下部干重均显著高于不接种对照,分别增加了60%和58%。与不接种对照相比,接种F. mosseae和G. versiforme处理的百合根尖数、根系长度、分叉数、表面积和根系体积分别比对照增加123%、128%、182%、118%和232%。切花瓶插期间,接种AM真菌处理的百合切花水分平衡值和鲜重变化率显著高于对照处理;乙烯释放速率和呼吸速率显著低于对照,瓶插5d时达到乙烯峰值5.4μL/*g·h (FW)+,共同接种F. mosseae和G. versiforme处理的百合切花乙烯释放速率比对照降低30%;呼吸速率则在瓶插1d时达到峰值0.7μL/*g·h (FW)+,共同接种比对照降低37%;百合花瓣内超氧化物歧化酶(SOD)活性、过氧化氢酶(CAT)活性、可溶性糖含量和可溶性蛋白含量比对照分别提高19%、32%、52%和26%。百合花瓣内N、P、K、Mg、Ca、Fe和Zn含量均显著高于不接种对照处理,Mn和Cr含量则低于对照;共同接种处理的百合瓶插寿命增加了3d,最佳观赏时间比对照延长2d。结论认为共同接种F. mosseae和G. versiforme处理更加有效地改善切花花枝的水分平衡、营养状况与生理代谢,控制衰老激素的合成,从而延长百合切花的瓶插寿命和最佳观赏期。  相似文献   

5.
分根培养系统中AM真菌抑制大豆胞囊线虫病的效应   总被引:3,自引:0,他引:3       下载免费PDF全文
在分根培养系统中将大豆(Glycine max Merrill)幼苗接种丛枝菌根(Arbuscular mycorrhiza,AM)真菌摩西球囊霉(Glomus mosseae)、幼套球囊霉(G.etunicatum)或/和大豆胞囊线虫(SCN,Heterodera glycines)后,定期测定大豆根系中AM真菌及线虫侵染率、病情指数、过氧化物酶(POD)、苯丙氨酸解氨酶(PAL)、β-1,3-葡聚糖酶及几丁质酶活性的动态变化。结果表明,将大豆胞囊线虫与AM真菌接种于分根培养系统同一室或分别接种于不同室,AM真菌均能显著降低大豆病情指数和线虫侵染速率,且将二者接种同一室处理的效果大于不同室处理;与只接种SCN的处理相比,接种摩西球囊霉和幼套球囊霉3周后在同室接种SCN处理的根围土壤中胞囊数、根上胞囊数和根内线虫数分别降低了47.4%、58.9%、46.6%和50.5%、67.0%、57.5%,表明AM真菌能显著抑制线虫的发育;摩西球囊霉和幼套球囊霉在一定时间段内诱导了大豆根系过氧化物酶、苯丙氨酸解氨酶、β-1,3-葡聚糖酶及几丁质酶的活性。摩西球囊霉和幼套球囊霉能够诱导大豆植株抵抗大豆胞囊线虫的侵染,既能在同一室的根围局部与SCN竞争侵染位点,又能对不同室的根系诱导产生防御性酶,推测摩西球囊霉和幼套球囊霉拮抗SCN的效应既是局部的,也是系统性的。  相似文献   

6.
喀斯特土壤上香樟幼苗接种不同AM真菌后的耐旱性效应   总被引:2,自引:0,他引:2  
为探索喀斯特土壤适生植物香樟幼苗在接种不同AM真菌后的耐旱适应性,进行了香樟幼苗接种幼套球囊霉(Glomus etunicatum)和层状球囊霉(Glomus lamellosum)后水分胁迫处理试验。结果表明:(1)接种AM真菌显著提高了香樟幼苗的生物量积累,AM促进植株生物量效应依次为中度>轻度>正常>重度,同一水分胁迫处理下生物量幼套球囊霉>层状球囊霉。(2)中度干旱下香樟幼苗菌根依赖性最大,幼套球囊霉接种植株的菌根依赖性较层状球囊霉大。(3)接种AM真菌显著提高了植株叶片可溶性糖、可溶性蛋白质和脯氨酸含量,并降低了丙二醛含量;在正常供水下植株叶片可溶性糖、可溶性蛋白质和脯氨酸含量层状球囊霉接种>幼套球囊霉接种>对照,干旱胁迫下表现为幼套球囊霉接种>层状球囊霉接种>对照;干旱胁迫下的幼套球囊霉接种植株丙二醛含量低于层状球囊霉接种植株。(4)总体上,可溶性糖与脯氨酸相关性极显著,可溶性蛋白质与丙二醛之间呈显著负相关性。幼套球囊霉接种香樟幼苗的耐旱性高于和层状球囊霉接种香樟幼苗。  相似文献   

7.
盐渍条件下AM真菌对大豆生长和离子含量的影响   总被引:6,自引:0,他引:6  
盐渍条件下研究了丛枝菌根(AM)真菌对大豆Glycine max植株生长和叶片离子含量的影响。结果表明,接种摩西球囊霉Glomus mosseae处理的叶片K+含量和K/Na比显著高于对照,而Na+含量无显著差异。G.mosseae显著增加了大豆植株生长量,这一效应随盐处理浓度的提高而增大。表明盐渍条件下AM真菌提高大豆抗盐性与其增加K+吸收和运输有关。  相似文献   

8.
通过盆栽试验研究了Na Cl胁迫下接种从枝菌根真菌(AMF)摩西球囊霉(Glomus mosseae)对红花(Carthamus tinctorius)生长及生理指标的影响。结果表明,接种摩西球囊霉能够显著提高了盐胁迫下红花地下部干重、地上部叶绿素含量,减少了丙二醛(MDA)的积累量。在盐处理下,接种AMF较不接种显著增加了红花叶片游离脯氨酸和可溶性糖的含量,提高了红花的总抗氧化能力。此外,接种AMF还能增强红花对矿质营养元素P、Mg的吸收。在高盐浓度(3‰)时,接菌较不接菌显著减少了红花对Na+的吸收,同时显著增加了红花叶片中K+的浓度。因此,摩西球囊霉能减少盐对叶绿素合成的干扰作用,从而提高叶绿素含量;同时通过提高渗透调节物质含量、增强矿质元素的吸收、减小膜脂过氧化程度以及降低高盐浓度时Na+的吸收来缓解红花的盐胁迫。  相似文献   

9.
高雁琳  李钧敏  闫明 《广西植物》2016,36(5):539-547
为改善采矿废弃物上植被生长状况,提高植物成活率,该研究采用盆栽试验法,以高丹草为材料,选用摩西球囊霉(Glomus mosseae,G.m)和地表球囊霉(G.versiforme,G.v)两种AM真菌,分别研究单接种和混合接种对粉煤灰(S1)、煤矸石(S2)和粉煤灰与煤矸石混合物(S3)三种煤矿废弃物基质上高丹草(Sorghum bicolor×S.sudanense)生长及叶绿素荧光的影响,并以正常沙土(S4)作为对照。结果表明:(1)4种基质上,3种接种处理均获得较高侵染率,在基质S1、S3和S4上均为接种摩西球囊霉对高丹草根系侵染率最高,分别为49.04%、57.40%、43.34%,在基质S2上,混合接种处理对高丹草根系侵染效果最好,达49.33%。(2)3种煤矿废弃物基质上高丹草根长、干重、叶绿素含量、F_v/F_o、q P和Yield显著降低。接种AM真菌显著提高了高丹草的生长和光合效率。与其他处理相比,在基质S1、S3和S4上,接种摩西球囊霉显著增加了根长、干重、叶绿素含量、F_v/F_o、q P和Yield,在基质S2上,接种地表球囊霉显著增加了根长、干重,接种地表球囊霉和摩西球囊霉+地表球囊霉(G.mv)处理间叶绿素荧光参数均无显著差异。这表明在煤矿废弃物基质的复合逆境中高丹草生长和光合作用显著受到抑制,AM真菌可通过提高高丹草叶绿素含量,改善叶片叶绿素荧光和光合作用,促进植物生长,来缓解该复合逆境对高丹草造成的伤害,增强其对煤矿废弃物不良环境的抗逆性,提高煤矿区植被恢复效果。接种摩西球囊霉对粉煤灰以及粉煤灰和煤矸石混合基质上高丹草的促进作用最佳,而接种地表球囊霉更适于煤矸石基质上高丹草的生长。  相似文献   

10.
 利用两种不同土壤研究了水分胁迫和接种AM真菌(摩西球囊霉(Glomus mosseae)和油蒿(Artemisia ordosica)根际土著AM真菌)对毛乌素沙地重要演替物种油蒿生长和抗旱性的影响。结果表明, 两种土壤中水分胁迫没有显著影响油蒿的植株形态和含水量, 但严重抑制了菌根侵染率。水分胁迫促使油蒿提高叶片保水能力, 抑制N、P在地上部的分配。在胁迫前期SOD活性较高, 而POD活性在后期较高。同一水分条件下接种AM真菌显著提高了AM真菌侵染率, 土壤中孢子数显著增多, 提高了植株分枝数并促进侧根发育, 显著提高根冠比和植株保水能力, 加强了根系对全磷、全氮的吸收。接种AM真菌的植株可溶性糖和丙二醛含量较低, 可溶性蛋白含量无显著变化, SOD和POD活性提高, 油蒿抗旱性加强。水分胁迫下在不同土壤中接种不同AM真菌对油蒿的促进效应差异较大, 接种土著AM真菌的效果优于摩西球囊霉单一接种。干旱导致菌根侵染率下降是宿主植物吸水能力下降的原因之一, 在植物生长前期接种AM真菌可以增强植物抵抗生长中后期环境干旱的能力。  相似文献   

11.
The phylogeny of Old World monkeys has remained unresolved in part because of a lack of resolution in the Cercopithecinae. Competing morphological hypotheses have had Allen's swamp monkey (Allenopithecus nigroviridis) and the talapoins (Miopithecus spp.) as basal branches of either the tribe Cercopithecini or the tribe Papionini. Previous molecular analyses have not adequately addressed the issue. To better understand the evolutionary history of these primates, we sequenced and subjected to phylogenetic analysis 3.1 kb of 2 loci (TSPY and SRY) from the non-recombining portion of the Y-chromosome. Individuals from the genera Allenopithecus, Miopithecus, Erythrocebus, Chlorocebus, and Cercopithecus were surveyed and their sequences compared with those previously published for the Papionini and Colobinae. The results suggest Allenopithecus and Miopithecus are more closely related to the Cercopithecini than Papionini. Our data also support the hypothesis that within the Cercopithecini, Erythrocebus and Chlorocebus share a close evolutionary relationship, distinct from the other members of the tribe.  相似文献   

12.
The inhibitory effect of nystatin and killer toxin on the growth of free and covalently-immobilizedSaccharomyces cerevisiae cells was studied. The resistance of immobilized cells to both agents was accompanied by increased amounts of phospholipids and sterols. The possible relationship between these changes in the membrane composition and the transduction of a signal across the cytoplasmic membrane is discussed.  相似文献   

13.
The formation of different structures in Drosophila depends on the combined activities of selector genes and signaling pathways. For instance, the antenna requires the selector gene homothorax, which distinguishes between the leg and the antenna and can specify distal antenna if expressed ectopically. Similarly, the eye is formed by a group of "eye-specifying" genes, among them eyeless, which can direct eye development ectopically. We report here the characterization of the hernandez and fernandez genes, expressed in the antennal and eye primordia of the eye-antenna imaginal disc. The predicted proteins encoded by these two genes have 27% common amino acids and include a Pipsqueak domain. Reduced expression of either hernandez or fernandez mildly affects antenna and eye development, while the inactivation of both genes partially transforms distal antenna into leg. Ectopic expression of either of the two genes results in two different phenotypes: it can form distal antenna, activating genes like homothorax, spineless, and spalt, and it can promote eye development and activates eyeless. Reciprocally, eyeless can induce hernandez and fernandez expression, and homothorax and spineless can activate both hernandez and fernandez when ectopically expressed. The formation of eye by these genes seems to require Notch signaling, since the induction of ectopic eyes and the activation of eyeless by the hernandez gene are suppressed when the Notch function is compromised. Our results show that the hernandez and fernandez genes are required for antennal and eye development and are also able to specify eye or antenna ectopically.  相似文献   

14.
Amplified ribosomal DNA restriction enzyme analysis (ARDRA), pulsed field gel electrophoresis (PFGE) and ribotyping were used to differentiate among 24 strains of Brevibacterium linens, Brevibacterium casei and Brevibacterium epidermidis obtained from type culture collections or isolated from various smear ripened cheeses. ARDRA was applied to the 16S rDNA. B. linens was shown to be a quite heterogenic group with 2 to at least 4 copies of rrn operons per strain with aberrant nucleotide sequences. AccI gave genus specific restriction patterns and was used to separate Brevibacterium from Corynebacterium species. The expected species specificity of TaqI applied to B. linens type culture strains, but not to all strains isolated from cheese. By AvaI restriction, B. casei and B. linens were differentiated from B. epidermidis and the orange pigmented Arthrobacter casei, a new species of coryneform bacteria; by XmnI restriction, B. linens and B. epidermidis were differentiated from B. casei. One of 4 B. linens genotypes could not be distinguished from B. casei by this method. Here, the typical orange B. linens pigments were used for classification, which was confirmed by partial sequencing of the 16S rDNA.  相似文献   

15.
Resin glycosides are secondary metabolites exclusive to the convolvulaceous plants. In this study, crypthophilic acids A–C (13), the first resin glycosides occurring in another family (Scrophulariaceae), and the other constituents of Scrophularia cryptophila were examined for in vitro antiprotozoal and antimycobacterial potentials. Except for crypthophilic acid B (2), all tested compounds exhibited growth-inhibitory effect against Trypanosoma brucei rhodesiense, with l-tryptophan (6) and buddlejasaponin III (7) being the most potent ones (IC50's 4.1 and 9.7 μg/ml). In contrast, the activity towards Trypanosoma cruzi was poor, and only crypthophilic acid C (3), 6 and 7 were trypanocidal at concentrations above 40 μg/ml. With the exception of 2 and 6, all compounds were active against Leishmania donovani. Harpagide (4) and 3 emerged as the best leishmanicidal agents (IC50's 2.0 and 5.8 μg/ml). Only compounds 3, 6 and 7 showed antimalarial activity against Plasmodium falciparum with IC50 values of 4.2, 16.6 and 22.4 μg/ml. Overall the best and broadest spectrum activity was presented by compounds 3 and 7, as they inhibited all four parasitic protozoa. None of the isolates had significant activity against Mycobacterium tuberculosis (MICs >100 μg/ml) or were toxic towards mammalian (L6) cells. This is the first report of antiprotozoal activity for natural resin glycosides, as well as for harpagide (4), acetylharpagide (5), tryptophan (6) and buddlejasaponin III (7).  相似文献   

16.
Aza- and deazaanalogues of adenosine, including their 1-protonated forms (except for that of 1-deazaadenosine), were studied by computer computation to find a relationship between their molecular structures and substrate properties for the mammalian adenosine deaminase. The atomic charge distribution and maps of the electrostatic potential around their van der Waals molecular surface were calculated for these compounds using the ab initio STO-3G method. The conformational studies are carried out by the MM+ method of molecular mechanics. The mechanism that determines the substrate selectivity of mammalian adenosine deaminase is discussed.  相似文献   

17.
Pericarp structure was investigated in 158 species of the familiesLamiaceae andVerbenaceae. Data from 221 out of 262 genera ofLamiaceae s.l. and a few ofVerbenaceae s.str. were collected in a table. A cladistic analysis was performed on the basis of pericarp characters only. The abandonment of subfam.Pogostemonoideae as a taxonomic unit is considered. Examples of groups given additional support by similarities in pericarp characters are: (1) the gynobasic-styled labiates (subfamiliesPogostemonoideae, Lamioideae, Nepetoideae); (2) aLamioideae-Pogostemonoideae-group; (3)Nepetoideae; (4) aWestringia-Hemigenia-Hemiandra-Microcorys group (in subfam.Chloranthoideae); (5) aLepechinia-Chaunostoma-group (inNepetoideae); (6) aPrunella-Cleonia-group (inNepetoideae).  相似文献   

18.
Effects of carbon concentration and carbon to nitrogen (C:N) ratio on six biocontrol fungal strains are reported in this paper. All fungal strains had extensive growth on the media supplemented with 6–12 g l−1 carbon and C:N ratios from 10:1 to 80:1, and differed in nutrient requirements for sporulation. Except for the two strains of Paecilomyces lilacinus, all selected fungi attained the highest spore yields at a C:N ratio of 160:1 when the carbon concentration was 12 g l−1 for Metarhizium anisopliae SQZ-1-21, 6 g l−1 for M. anisopliae RS-4-1 and Trichoderma viride TV-1, and 8 g l−1 for Lecanicillium lecanii CA-1-G. The optimal conditions for P. lilacinus sporulation were 8 g l−1 carbon with a C:N ratio of 10:1 for M-14 and 12 g l−1 carbon with a C:N ratio of 20:1 for IPC-P, respectively. The results indicated that the influence of carbon concentration and C:N ratio on fungal growth and sporulation is strain dependent; therefore, consideration for the complexity of nutrient requirements is essential for improving yields of fungal biocontrol agents.  相似文献   

19.
K. Schick  N. Toth 《Human Evolution》2000,15(1-2):121-128
Around two-and-a-half million years ago, some hominid populations in Africa began to modify stones and bones in a manner that can be recognized by prehistorians as artifacts, and, by definition, produced the earliest identifiable archaeological record. It is likely that earlier hominid groups also may have had relatively rich tool-using behavioral repertoires similar to that seen in modern chimpanzees (McGrew, 1992), such tools may have been made of perishable materials or minimally modified and thus difficult to identify. This review will focus on the earliest archaeological traces and the spread of hominids out of Africa and into Eurasia.  相似文献   

20.
Spatial and temporal patterns of morel fruiting   总被引:2,自引:0,他引:2  
The biotic and abiotic factors conditioning morel fruit body production are incompletely known. We examined spatial and temporal patterns of Morchella esculenta fruiting over five years in a wooded site in Missouri, USA. Fruiting onset was inversely correlated with spring air and soil temperatures, whereas abundance was positively correlated with rain events (>10 mm) during the 30 d preceding fruiting. The two years with the greatest fruiting had the shortest fruiting seasons (6–7 d). Fruiting season length was positively correlated with soil warming, suggesting that a narrow range of optimum soil temperatures favour the explosive production of fruit bodies. All woody stems of at least 1 cm diam were mapped and stem diameter and crown condition were noted. Morel fruit bodies were significantly closer to stems of Carya spp., Tilia americana and Ulmus americana than predicted by the frequencies of these woody species or their contribution to the total basal area on the site. Although intra-annual clustering of fruit bodies was often observed, inter-annual clustering was not. The spatial pattern of M. esculenta fruiting appears to be associated with vegetation pattern, whereas the onset and abundance of fruiting are determined by the interaction of spring temperatures with availability of supporting precipitation.  相似文献   

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