A LAMP-assay-based specific microbiota analysis reveals community dynamics and potential interactions of 13 major soybean root pathogens

Soybean root diseases are associated with numerous fungal and oomycete pathogens;however,the community dynamics and interactions of these pathogens are largely unknown.We performed 13 loop-mediated isothermal amplification(LAMP)assays that targeted specific soybean root pathogens,and traditional isolation assays.A total of 159 samples were collected from three locations in the Huang-Huai-Hai region of China at three soybean growth stages(30,60,and 90 days after planting)in 2016.In LAMP results,we found that pathogen communities differed slightly among locations,but changed dramatically between soybean growth stages.Phytophthora sojae,Rhizoctonia solani,and Fusarium oxysporum were most frequently detected at the early stage,whereas Phomopsis longicolla,Fusarium equiseti,and Fusarium virguliforme were most common in the later stages.Most samples(86%)contained two to six pathogen species.Interestingly,the less detectable species tended to exist in the samples containing more detected species,and some pathogens preferentially co-occurred in diseased tissue,including P.sojae–R.solani–F.oxysporum and F.virguliforme–Calonectria ilicicola,implying potential interactions during infection.The LAMP detection results were confirmed by traditional isolation methods.The isolated strains exhibited different virulence to soybean,further implying a beneficial interaction among some pathogens.

Model Construction and Visualization Simulation of Soybean Root System

This paper mainly focused on the growth law, model construction and visualization of a soybean root. A pot experiment was conducted in the laboratory to collect root system data and measure soybean root＇s length, diameter and number by excavating in different periods. On the basis of an in-depth analysis of root structure geometry, we analyzed the collected experimental data with logistic equations, and got the growth of soybean root equation, according to its morphological structure characteristics of self-similarity, and discussed the virtual modeling method on the soybean root based on L-system and in Visual c＋＋ Using OpenGL technology to achieve a soybean root system topology model and visualization simulation.

Effect of plough pans on the growth of soybean roots in the black-soil region of northeastern China

Plough pans （PPs）, common in the black-soil region of northeastern China, have significant effects on the root systems of crops. We conducted a field experiment to study the distribution of soybean roots under the influence of PPs. The soybean roots showed compensatory growth above the PP, with higher root length and weight in soil with a PP compared to those without a PP. Roots were heavier and longer in the 15-75 cm soil layer without a PP than with a PP. Soil porosity was lower in the PP and the soil below the PP, which likely decreased the oxygen content of the soil and induced more growth of roots above the PP. The PP is also likely to decrease infiltration of rain-water and hinder the migration of nitrate downward, which in turn increased the density and length of soybean root hairs, which, hence, promoted growth.

Impact of long-term chemical fertilizer and organic amendment to Fusarium root rot of soybean

Soil suppressiveness to Fusarium root rot of soybean had been observed in a black soil field after a long-term fertilization with nitrogen(N)and phosphorus(P)fertilizer combined with pig manure as organic amendment(NPM),rather than that with only nitrogen and phosphorus fertilizer(NP)or no fertilizer(NF).To determine the microbial role on this suppressiveness,fungal and bacterial community characteristics in NPM,NP and NF treatments were investigated by q PCR and DGGE.Compared with the similar bacterial community characteristics among 3 treatments,fungal community,especially Fusarium population size and community composition in NPM treatment were different with those of NP and NF groups.Based on the isolation and pathogenicity test,pathogenic F.oxysporum,F.graminearum,F.verticillioide and F.lateritium absolutely dominated Fusarium community in NF and NP groups.Nonpathogenic F.avenaceum,F.equiseti,F.culmorum,F.redolens,F.solani and F.tricinctum dominated Fusarium community in NPM group.Isolation rate of pathogenic Fusarium in NPM reduced from 100%to 38%in NF.These results suggested that the dominance of soil non-pathogenic Fusarium population induced by organic amendment might play an important role on suppressing Fusarium root rot in the tested field.

Overexpression of GmBIN2,a soybean glycogen synthase kinase 3 gene,enhances tolerance to salt and drought in transgenic Arabidopsis and soybean hairy roots

Glycogen synthase kinase 3 （GSK3） is a kind of sedne/threonine kinase widely found in eukaryotes. Many plant GSK3 kinases play important roles in regulating stress responses. This study investigated BRASSINOSTEROID-INSENSITIVE 2 （GmBIN2） gene, a member of the GSK3 protein kinase family in soybean and an orthologue of Arabidopsis BIN2/AtSK21. GmBIN2 expression was increased by salt and drought stresses, but was not significantly affected by the ABA treatment. To examine the function of GrnBIN2, transgenic Arabidopsis and transgenic soybean hairy roots were generated. Overexpression of GmBIN2 in Arabidopsis resulted in increased germination rate and root length compared with wild-type plants under salt and mannitol treatments. Overexpression of GmBIN2 increased cellular Ca2~ content and reduced Na~ content, enhancing salt tolerance in transgenic Arabidopsis plants. In the soybean hairy root assay, overexpression of GmBIN2 in transgenic roots also showed significantly higher relative root growth rate than the control when subjected to salt and mannitol treatments. Measurement of physiological indicators, including proline content, superoxide dismutase （SOD） activity, and relative electrical conductivity, supported this conclusion. Furthermore, we also found that GmBIN2 could up-regulate the expression of some stress-related genes in transgenic Arabidopsis and soybean hairy roots. Overall, these results indicated that GmBIN2 improved tolerance to salt and drought in transgenic Arabidopsis and soybean hairy roots.

Gm NAC15 overexpression in hairy roots enhances salt tolerance in soybean

The NAC （NAM, ATAF1/2 and CUC2） transcription factor family plays a key role in plant development and responses to abiotic stress. GmNAC15 （Glyma 15g40510.1）, a member of the NAC transcription factor family in soybean, was functionally characterized, especially with regard to its role in salt tolerance. In the present study, qRT-PCR （quantitative reverse transcription PCR） analysis indicated that GmNAC15 was induced by salt, drought, low temperature stress, and ABA treatment in roots and leaves. GmNAC15 overexpression in soybean （Glycine max） hairy roots enhanced salt tolerance. Transgenic hairy roots improved the survival of wild leaves; however, overexpression of GmNAC15 in hairy root couldn＇t influnce the expression level of GmNAC15 in leaf. GmNAC15 regulates the expression levels of genes responsive to salt stress. Altogether, these results provide experimental evidence of the positive effect of GmNAC15 on salt tolerance in soybean and the potential application of genetic manipulation to enhance the salt tolerance of important crops.

Soybean hairy roots produced in vitro by Agrobacterium rhizogenes-mediated transformation

Soybean is one of the world's most important oil and protein crops. Efficient transformation is a key factor for the improvement of soybean by genetic modification. We describe an optimized protocol for the Agrobacterium rhizogenes-mediated transformation of soybean and the induction of hairy root development in vitro. Cotyledons with 0.5-cm hypocotyls were cut from 5-day-old seedlings and used as explants. After infection and co-cultivation,hairy roots were produced in induction culture medium after 10–12 days. Using this method, 90%–99% of the infected explants of five different cultivars produced hairy roots within one month. Observations using reporter constructs showed that 30%–60% of the hairy roots induced were transformed. Based on high transformation efficiency and short transformation period, this method represents an efficient and rapid platform for study of soybean gene function.

Resistance Evaluation of Soybean Germplasm from Huanghuai Region to Phytophthora Root Rot

The aim of the study was to establish a set of differential strains and to identify soybean resistant genes to Phytophthora root rot and then to apply those strains for analysis of the resistant genes Rps1a,Rps1c,and Rps1k that soybean cultivars or lines may carry.Virulence formula of 125 Phytophthora sojae isolates were determined using the hypocotyls inoculation technique,the strains,which includ 6 isolates with different virulence formulas,were applied to identify the resistance of 55 soybean cultivars or lines and resistant genes were analyzed using the gene postulating procedure.Eighteen reaction types occurred in 55 cultivars or lines and results of gene postulation indicated that 2 cultivars or lines probably carried gene Rps1c and no cultivar may carry genes Rps1a or Rps1k.A few of soybean cultivars or lines from Huanghuai Region carry Rps genes Rps1a,Rps1c and Rps1k and tend to infect by P.sojae,so resistant cultivars or lines need to be bred and popularized actively.

Evolution of Root Characters of Soybean Varieties Developed in Different Years

It was studied that the evolution of root characteristics among 42 soybean varieties developed in Heilongjiang and Jilin Province in different years. The results showed that there were differences on the root characteristics among soybean varieties. From 1950s to 1990s, root fresh weight, root volume, root surface, root dry weight, lateral root length of main root characters tendedly increased with the variable development

Hypoxia-Responsive Root Hydraulic Conductivity Influences Soybean Cultivar-Specific Waterlogging Tolerance

Excess soil moisture induces hypoxic conditions and causes waterlogging injury in soybean [Glycine max (L.) Merr.]. This study investigated the mechanism underlying the development of waterlogging injury. Nine Japanese soybean cultivars with varying degrees of waterlogging tolerance were grown in a hydroponic system for 14 days under hypoxic conditions. Shoot and root biomasses and root hydraulic conductivity were measured at an early vegetative stage for plants under control and hypoxic conditions. Root morphological traits and intramembrane aquaporin proteins were also analyzed. The tolerance of each cultivar to field waterlogging was based on biomass changes induced by the hypoxia treatment. Root hydraulic conductivity responses to hypoxia were associated with changes in total dry weight, leaf dry weight, and leaf area. The effects of hypoxic conditions on root hydraulic conductivity were also represented by the changes in root morphology, such as total root length, thick-root length, and number of root tips. Additionally, a 32.3 kDa aquaporin-like protein seemed to regulate root hydraulic conductivity. Our results from a hydroponic culture suggest that the soybean cultivar-specific responses to hypoxic conditions in the rhizosphere reflect fluctuations in hydraulic conductivity related to root morphological or qualitative changes.

Conidia of one Fusarium solani isolate from a soybean-production field enable to be virulent to soybean and make soybean seedlings wilted

Fusarium is usually thought to cause soybean root rot, which results in a large quantity of annual yield loss in soybean production, by its secretions including Fusarium toxins and cell wall degrading enzymes, but not by the conidia themselves that do not underlie any virulence so far. Here we report that the conidia of one Fusarium solani isolate are able to be virulent to soybean and make soybean seedlings wilted alone. We isolated them from the wilted plants in a soybean-production field and molecularly identified 17 Fusarium isolates through phylogenetic analysis. Of them, except for one isolate that showed diversity of virulence to different soybeans （virulent to one soybean whereas avirulent to another soybean）, the others were all virulent to the two tested soybeans： both conidia cultures and secretions could make soybean seedlings wilted at 5 days post infection, and their virulence had dosage effects that only conidia cultures of at least 5×10^6 conidia mL-1 could show virulence to soybean; however, the sole conidia of the F. solani isolate #4 also exhibited virulence to soybean and could make soybean seedlings wilted. Finally, we developed the specific cleaved amplified polymorphic sequences （CAPS） markers to easily differentiate Fusarium isolates. The isolate #4 in this work will likely be used to investigate the new mechanism of virulence of Fusarium to soybean.

Resistance identification of bivalent fungi-resistant genes transformed soybean to Phytophthora sojae

Soybean is one of the most important sources of edible oil and proteins in the world. However, it suffers from many kinds of fungal diseases which is a major limiting factor in soybean production. The fungal disease can be effectively controlled by breeding plant cultivars with genetic transformation. In this study, the resistance to Phytophthora sojae of five bivalent transgenic soybean lines was identified using the hypocotyls inoculation technique. The lines were the T2 of the transgenic soybean which were transformed with kidney bean chitinase gene and barley ribosome inactivating protein gene, and were positive by Southern Blot analysis. The resistance difference was studied through comparing the death percentage of transgenic soybean with the control. The results showed that four lines were more resistant to P sojae, whereas other one had no significant difference in comparison with the control. These transgenic soybean lines with enhanced resistance to P sojae will be useful in soybean resistance breeding.

Differences between soybean genotypes in physiological response to sequential soil drying and rewetting

Soybean genotypes show diverse physiological responses to drought, but specific physiological traits that can be used to evaluate drought tolerance have not been identified. In the present study we investigated physiological traits of soybean genotypes under progressive soil drying and rewetting, using a treatment mimicking field conditions.After a preliminary study with eight soybean genotypes, two drought-tolerant genotypes and one susceptible genotype were grown in the greenhouse and subjected to water restriction. Leaf expansion rate, gas exchange, water relation parameters, total chlorophyll(Chl), proline contents of leaves, and root xylem p H were monitored in a time course, and plant growth and root traits were measured at the end of the stress cycle. Drought-tolerant genotypes maintained higher leaf expansion rate, net photosynthetic rate(Pn), Chl content,instantaneous water use efficiency(WUEi), % relative water content(RWC), water potential(ψw), and turgor potential(ψp) during progressive soil drying and subsequent rewetting than the susceptible genotypes. By contrast, stomatal conductance(gs) and transpiration rate(Tr)of tolerant genotypes declined faster owing to dehydration and recovered more sharply after rehydration than the same parameters in susceptible ones. Water stress caused a significant increase in leaf proline level and root xylem sap p H of both genotypes but tolerant genotypes recovered to pre-stress levels more quickly after rehydration. Tolerant genotypes also produced longer roots with higher dry mass than susceptible genotypes. We conclude that rapid perception and adjustment in response to soil drying and rewetting as well as the maintenance of relatively high Pn, %RWC, and root growth constitute the mechanisms by which drought-tolerant soybean genotypes cope with water stress.

大豆镰刀菌根腐病拮抗菌的筛选及生防效果

【目的】镰刀菌根腐病是世界范围内大豆生产上最具破坏性的土传病害之一,为获得对禾谷镰刀菌Fusarium graminearum具有较好拮抗效果的生防菌株。【方法】从健康大豆根际土壤分离细菌,平板对峙法筛选拮抗菌株,通过形态观察、生理生化特性、胞外酶活性及促生特性对菌株进行鉴定分析,采用盆栽试验进一步测定菌株生防及促生效果。【结果】筛选出的4株芽孢杆菌和1株假单胞杆菌对F.graminearum的抑菌率均达到60%以上,对F.oxysporum,F.solani,F.longifundum以及F.equiseti也均有一定的抑制作用,其发酵液及挥发代谢物均会影响F.graminearum的生长。5株拮抗菌具有产蛋白酶、纤维素酶以及葡聚糖酶的活性,解磷、解钾、固氮以及产铁载体的能力。综合以上测试结果,菌株20-8具有较强的抑菌及大豆促生效果。根据形态特征及16S rRNA序列分析,将菌株20-8鉴定为暹罗芽孢杆菌(Bacillus siamensis)。该菌株的发酵上清液可以破坏F.graminearum菌丝体结构,无细胞发酵上清液可以显著抑制孢子萌发。室内盆栽防效测定结果表明,20-8的稀释发酵液对F.graminearum引起大豆根腐病的防效可达46.08%,并且促进大豆植株生长。【结论】筛选鉴定的暹罗芽孢杆菌20-8具有解磷、解钾、固氮以及产铁载体及多种胞外酶功能,其稀释发酵液具有较强的抗真菌活性及大豆促生能力,菌株20-8可以用于防治F.graminearum引起的大豆根腐病。

GmYUC12a对大豆侧根发育和抗旱性的调控作用

侧根对作物抗旱具有重要作用,明确Gm YUC12a在大豆侧根生长发育和抗旱中的作用,为探究大豆侧根发育响应干旱的作用机制奠定基础。本文首先分析了徐豆18和徐9302在5%(m/V)聚乙二醇6000模拟干旱胁迫下根系形态特征、根系内源生长素(IAA)含量、IAA合成和信号转导相关基因表达量以及叶片净光合速率(P_(n))和相对含水量(RWC)的变化。干旱胁迫下根系IAA含量、Gm YUC12a、Gm TIR1A和Gm AFB3A表达量变化趋势与侧根数、侧根长、总根长一致,徐9302各项指标均较对照增加,徐豆18则相反,因而徐豆18叶片P_(n)和RWC下降幅度较大。通过发根农杆菌侵染徐9302下胚轴,获得转基因毛状根嵌合植株。干旱胁迫下Gm YUC12a过表达毛状根嵌合体根系IAA含量、Gm TIR1A、Gm AFB3A表达水平、叶片P_(n)和RWC以及侧根数量和长度显著高于空载毛状根嵌合体。与之不同,干旱胁迫下Gm TIR1A、Gm AFB3A过表达毛状根嵌合体侧根数量和长度较空载毛状根复合体无变化,而GmTIR1A和Gm AFB3A同时过表达毛状根嵌合体侧根数量和长度显著增加。可见,干旱胁迫下Gm YUC12a诱导GmTIR1A和Gm AFB3A表达,进而Gm TIR1A和Gm AFB3A协同调控侧根生长发育,提高植株抗旱性。

干旱胁迫下GmYUC12a对大豆侧根蔗糖代谢的影响

黄素单加氧酶编码基因(YUCs)是依赖于色氨酸生长素合成途径中的关键因子。为了明确GmYUC12a在大豆侧根蔗糖代谢响应干旱胁迫中的作用,试验以大豆稳定品系徐9302为材料,通过发根农杆菌侵染子叶下胚轴,获得过量表达发状根复合体植株,测定5%PEG 6000(M/V)模拟干旱胁迫下GmYUC12a过量表达发状根分生出的侧根形态特征、可溶性糖和蔗糖含量及蔗糖代谢相关基因表达量。结果发现,干旱胁迫下GmYUC12a过量表达显著增加侧根长度和干重,表明GmYUC12a可促进干旱胁迫下侧根的生长发育过程。GmYUC12a过量表达能够引起干旱胁迫下侧根GmSUC2、GmCWINV1和GmMST2表达上调,提高蔗糖卸载能力;促使GmSPS1、GmSPS2和GmSuSy1表达上调,增强蔗糖循环能力,促进侧根蔗糖和可溶性糖积累。综上,干旱胁迫通过上调GmYUC12a表达,调控侧根蔗糖卸载和循环过程,维持侧根生长。本研究结果揭示了干旱胁迫下GmYUC12a调控侧根发育的作用机制,为深入研究生长素合成在大豆干旱响应中的功能提供了参考。

Soybean Nodulation and Plant Response to Nitrogen and Sulfur Fertilization in the Northern US

Soybean [Glycine max (L.) Merrill] seed yields in the northern United States may increase with the application of fertilizers;however Nitrogen (N) may decrease root nodulation. This study was conducted to understand the impact of N and sulfur (S) fertilization on soybean nodulation, plant, shoot and root biomass. Two cultivars were planted in experiments across ten site-years during 2015-2016. Plant observations took place at the V4 and R4 soybean growth stages. There were 41% more nodules per plant at R4 compared to V4 (38.3 vs 27.2 nodules, respectively). Cultivars responded differently to N and S fertilizer. The nodules per plant between the cultivars (30.3 vs 24.4) were different as well as the percent medium and large-sized nodules, which indicates the need to evaluate additional genotypes. Adding N decreased root nodulation (from 31.8 to 23.7 nodules per plant) and decreased nodule size but had no effect on plant, shoot or root mass. Averaged across N rates total plant mass was 2.26 and 11.36 g per plant at V4 and R4, respectively. Shoot mass, average across N rates was 1.77 and 9.65 g per plant at V4 and R4, respectively, and root mass, average across N rates was 0.49 and 1.71 g per plant at V4 and R4, respectively. Sulfur did not have an effect on nodules per plant but increased the percent medium size nodules at the R4 observation. There was no N by S interaction observed for nodule number, size of the nodules, and plant, root and shoot mass. As cultivars differed in their nodulation response to N and S, additional research would be helpful to screen other cultivars.

Relationships between Microsclerotia Content and Hyperspectral Reflectance Data in Soybean Tissue Infected by Macrophomina phaseolina

Alternative methods are needed to assess the severity of charcoal rot disease [Macrophomina phaseolina (Tassi) Goid] in soybean [Glycine max (L.)] plant tissue. The objective of this study was to define the relationship between light reflectance properties and microsclerotia content of soybean stem and root tissue. Understanding that relationship could lead to using spectral reflectance data as a tool to assess the severity of charcoal rot disease in soybean plants, thus reducing human bias associated with qualitative analysis of soybean plant tissue and cost and time issues connected with quantitative analysis. Hyperspectral reflectance measurements (400-2490 nm) were obtained with a non-imaging spectroradiometer of non-diseased and charcoal rot diseased ground stem and root tissue samples of six soybean genotypes (“Clark”, “LD00-3309”, “LG03- 4561-14”, “LG03-4561-19”, “Saline”, and “Y227-1”). Relationships between the reflectance measurements and tissue microsclerotia content were evaluated with Spearman correlation (rs) analysis (p < 0.05). Moderate (rs = ±0.40 to ±0.59), strong (rs = ±0.60 to ±0.79), and very strong (rs = ±0.80 to ±1.00) negative and positive statistically significant (p < 0.05) monotonic relationships were observed between tissue spectral reflectance values and tissue microsclerotia content. Near-infrared and shortwave-infrared wavelengths had the best relationships with microsclerotia content in the ground tissue samples, with consistent results obtained with near-infrared wavelengths in that decreases in near-infrared spectral reflectance values were associated with increases in microsclerotia content in the stem and root tissue of the soybean plants. The findings of this study provided evidence that relationships exist between tissue spectral reflectance and tissue microsclerotia content of soybean plants, supporting spectral reflectance data as a means for assessing variation of microsclerotia content in soybean plants. Future research should focus on the modelling capabilities of the selected wavelengths and on the feasibility of using these wavelengths in machine learning algorithms to differentiate non-diseased from charcoal rot diseased tissue.

大豆根茎过渡区弯曲突变体M_(rstz)的鉴定与基因定位

植物根茎过渡区(root-stem transition zone,RSTZ)将根和茎相互连接,其发育形态决定了大豆的地上部株型和抗倒伏潜力。本研究通过EMS诱变获得一个RSTZ弯曲或旋转的大豆突变体M_(rstz),其形态特征能够稳定遗传,是探究大豆茎秆发育规律的特异材料。将该突变体和栽培大豆中黄13杂交构建重组自交系群体,对群体中直立和弯曲型后代的RSTZ进行解剖结构比较,发现弯曲型株系比直立型株系的维管形成层较宽、次生木质部细胞层数较多、细胞形状不规则,表明维管组织分化可能是导致RSTZ形态发生差异的重要因素之一。进一步对化学组分测定发现,木质素和粗纤维含量越高越不易弯曲。选取RIL群体中弯曲型和直立型2种极端株系进行BSA-Seq,采用SNP-index和InDel-index关联分析方法鉴定到调控RSTZ形态的关联区域Chr19:43030943~45849854,该区间共含有319个基因。结合生物信息学分析、基因注释信息和表达丰度分析筛选到7个候选基因,分别为Glyma.19G170200、Glyma.19G201500、Glyma.19G187800、Glyma.19G178200、Glyma.19G197000、Glyma.19G179100、Glyma.19G196900。其中,Glyma.19G187800、Glyma.19G178200和Glyma.19G196900在大豆驯化中潜在影响RSTZ形态建成。本研究为解析大豆RSTZ组织形成及其遗传基础提供了材料基础,并为挖掘调控大豆茎秆发育基因提供新的见解。

不同时期水分胁迫对大豆根系特性及产量的影响

为揭示抗旱基因型大豆根系应答干旱胁迫的响应机制,于2020年在辽宁省沈阳市开展盆栽试验,以抗旱基因型和干旱敏感型大豆为试材,探讨干旱(W1,土壤含水量为田间持水量的50%)、轻度干旱(W2,土壤含水量为田间持水量的60%)、适宜水分(W3,土壤含水量为田间持水量的80%)对大豆根系特性和产量的影响。结果表明:同一时期控水,W1和W2条件下,抗旱品种“辽豆14”的总根长和根表面积优于干旱敏感型品种“辽豆21”。一般情况下,平均根直径为0~0.5 mm的根长比例表现为:“辽豆21”>“辽豆14”,而平均根直径为0.5~1.0 mm的根长比例却表现为:“辽豆14”>“辽豆21”。随着生育进程的推进,大豆地上部鲜重、地上部干重、根干重逐渐累积,鼓粒期达最大值;一般情况下,同一控水时期,随着土壤含水量的增加,同一品种地上部鲜重、根鲜重、地上部干重、根干重值逐渐增加,但抗旱品种“辽豆14”的值一般优于干旱敏感型品种“辽豆21”。随着生育进程的推进,根冠比鲜重和根冠比干重值降低;土壤干旱提高了植株的根冠比。各个时期控水,品种和水分均显著影响了大豆的单株产量,干旱(W1)和轻度干旱(W2)均降低了2品种的单株产量,但抗旱品种“辽豆14”引起产量下降的值更低。干旱胁迫条件下,抗旱基因型大豆在总根长、根表面积、地上部干重、根干重、产量以及平均根直径为0.5~1.0 mm的根长比例方面,表现出优于干旱敏感型品种的优势。