Bacterial diversity of the rhizosphere of maize (Zea mays) grown in tropical soil studied by temperature gradient gel electrophoresis

The bacterial diversity and population dynamics in the rhizosphere of two maize cultivars (Nitroflint and Nitrodent) grown in tropical soils was studied, by traditional cultivation techniques and 16S rRNA gene-based molecular analysis of DNA directly extracted from soil and rhizosphere samples. Rhizosphere and soil samples were taken at three different plant growth stages. Total aerobic bacterial counts were determined. Fingerprints of the most dominant bacterial population were generated by TGGE separation of 16S rRNA gene fragments amplified from total community DNA using eubacterial specific primers. To reduce the complexity of TGGE fingerprints or to analyse less abundant populations, primers specific for different phylogenetic groups have been used. A comparison of the cfu obtained for rhizosphere of both cultivars indicated significant differences only for rhizosphere and soil samples taken 40 days after sowing. However, a comparison of TGGE patterns indicated that the composition of the bacterial community analysed at different plant growth stages for both cultivars was similar. A comparison of -, -proteobacterial and actinomycete TGGE patterns of both cultivars confirmed this observation. The eubacterial TGGE profiles reflected strong seasonal population shifts in the bacterial rhizosphere community of both maize cultivars which could be also observed in the TGGE patterns of - and -proteobacteria and to a lesser extent for actinomycetes. The rhizosphere effect was much more pronounced for young roots compared to samples taken from mature maize plants. The rhizosphere fingerprints showed a reduced complexity for young plants with up to five dominating bands while for mature plants patterns similar to those of soil were observed. Sequencing of dominant clones indicated that the dominant population found at all plant growth stages can be assigned to Arthrobacter populations.     

Zinc deficiency and pollen fertility in maize (Zea mays)

Zinc deficiency decreased pollen viability in maize (Zea mays L. cv. G2) grown in sand culture. On restoring normal zinc supply to zinc-deficient plants before the pollen mother cell stage of anther development, the vegetative yield of plants and pollen fertility could be recovered to a large extent, but the recovery treatment was not effective when given after the release of microspores from the tetrads. If zinc deficiency was induced prior to microsporogenesis it did not significantly affect vegetative yield and ovule fertility, but decreased the fertility of pollen grains, even of those which visibly appeared normal. If the deficiency was induced after the release of microspores from the tetrads, not only vegetative yield and ovule fertility but pollen fertility also remained unaffected.     

Isolation and characterization of trypsin inhibitors from cowpea (Vigna unguiculata)

The trypsin inhibitor fraction from cowpea (Vigna unguiculata) has been purified and characterized. Although the total trypsin inhibitor as purified by affinity chromatography on immobilised trypsin was shown to be heterogeneous by gel electrophoresis and isoelectric focusing as well as by function, it was relatively homogeneous in MW (ca 17 000) on gel filtration. The total trypsin inhibitor was divided into inhibitors active against trypsin only and active against trypsin and chymotrypsin by affinity chromatography on immobilised chymotrypsin. The ‘trypsin-only’ inhibitor was the major component of the total trypsin inhibitor. It was shown by isoelectric focusing and gel electrophoresis to contain several isoinhibitors. Determination of the combining weight of this inhibitor and investigation of the complexes formed with trypsin by gel filtration indicated the presence of two protease binding sites per inhibitor molecule. The chymotrypsin/trypsin inhibitor was also shown to be composed of several isoinhibitors. On the basis of gel electrophoresis and gel filtration in dissociating and non-dissociating media both inhibitors were considered to be dimeric molecules with the subunits linked by disulphide bonds; this implies that the ‘trypsin-only’ inhibitor has one binding site per subunit.     

Differences on Pb accumulation among plant tissues of 25 varieties of maize (Zea mays)

Pollution of agricultural land by heavy metals has imposed an increasingly serious risk to environmental and human health in recent years.Heavy metal pollutants may enter the human food chain through agricultural products and groundwater from the polluted soils.Progress has been made in the past decade on phytoremediation,a safe and inexpensive approach to remove contaminants from soil and water using plants.However,in most cases,agricultural land in China cannot afford to grow phytoremediator plants instead of growing crops due to food supply for the great population.Therefore,new and effective methods to decrease the risk of heavy metal pollution in crops and to clean the contaminated soils are urgently needed.If we can find crop germplasms (including species and varieties) that accumulate heavy metals in their edible parts,such as the leaves of vegetables or grains of cereals,at a level low enough for safe consumption,then we can grow these selected species or varieties in the lands contaminated or potentially contaminated by heavy metals.If we can find crop germplasms that take in low concentrations of heavy metals in their edible parts and high content of the metals in their inedible parts,then we can use these selected species or varieties for soil remediation.In this study,the feasibility of the method is assessed by analyzing Pb concentrations in edible and inedible parts of 25 varieties of maize (Zea mays) grown in Pb-contaminated soils.The soil concentrations of Pb were 595.55 mg/kg in the high Pb exposed treatment and 195.55 mg/kg in the control.The results showed that the Pb concentrations in different tissues were in the order of root ＞ shoot ≌ leaf＞ grain.Compared with the control,the Pb concentrations in root,shoot and leaf were greatly increased under the high Pb exposed condition,while the increments of Pb concentration in grain were relatively lower.Under the high Pb exposure,the grain Pb concentrations of 12 varieties exceeded the maximal Pb limitation of the National Food Hygiene Standard of China (NFHSC) and were inedible.This indicates that there is a high Pb pollution risk for maize grown on Pb polluted sites.Although 22 of the 25 tested varieties had harvest loss under the highly Pb stressed condition,ranging from 0.86%-38.7% of the grain biomass acquired at the control,the average harvest loss of all the tested varieties was only 12.6%,which is usually imperceptible in normal farming practices.Therefore the risk of Pb pollution in maize products cannot be promptly noticed and prevented based only on the outcome of the harvest.However,we did find that 13 of the 25 tested varieties had grain Pb concentrations lower than the limitation of the NFHSC.It is,therefore,possible to reduce the pollution risk if these favorable varieties are used for maize production in Pb-contaminated or potentially contaminated agricultural lands.Pb concentrations in vegetative tissues (root,stem and leaf) were significantly correlated with each other,while Pb concentrations of each vegetative tissue were not significantly correlated with that of grain.Among the 25 tested varieties,some varieties had Pb concentrations in grain lower than (No.1-3 and No.6) or slightly above (No.4) the limitation of the NFHSC,while their Pb concentrations in the vegetative tissues were among the highest.When excluding these varieties,correlations between the Pb concentrations of grain and those of vegetative tissues of the rest of the tested varieties became highly significant.In addition,variety No.1 had the lowest harvest loss under high Pb exposed,and the highest Pb accumulation in vegetative tissues (51.69 mg/plant,12 times as much as in the control).Similar features were also observed in varieties No.2,No.3 and No.6,which absorbed Pb for 36-42 mg/plant under high Pb exposed.We recommend these varieties of maize to be used for bioremediation of Pb contaminated soil and crop production at the same time.     

Distribution of the glutamine synthetase isozyme GSp1 in maize (Zea mays)

In maize (Zea mays L.), GSp1, the predominant GS isozyme of the developing kernel, is abundant in the pedicel and pericarp, but absent from the endosperm and embryo. Determinations of GSp1 tissue distribution in vegetative tissues have been limited thus far to root and leaves, where the isozyme is absent. However, the promoter from the gene encoding GSp1 has been shown to drive reporter gene expression not only in the maternal seed-associated tissues in transgenic maize plants, but also in the anthers, husks and pollen (Muhitch et al. 2002, Plant Sci 163: 865-872). Here we report chromatographic evidence that GSp1 resides in immature tassels, dehiscing anthers, kernel glumes, ear husks, cobs and stalks of maize plants, but not in mature, shedding pollen grains. RNA blot analysis confirmed these biochemical data. In stalks, GSp1 increased in the later stages of ear development, suggesting that it plays a role in nitrogen remobilization during grain fill.     

土壤耕作及秸秆还田对夏玉米田杂草生物多样性的影响

于2008年夏玉米(Zea mays)生长期间,在连续5a秸秆全量还田的免耕、旋耕、耙耕、深松和常规耕作试验地中,研究了杂草总密度、优势杂草种类、生物多样性指数、杂草生物量和夏玉米产量。调查共记录杂草种类13种,秸秆全量还田时,免耕显著提高杂草的总密度;无秸秆还田时,常规耕作的杂草密度高于免耕、旋耕、耙耕和深松。秸秆全量还田后,免耕和深松条件下,杂草优势种为马唐和旱稗,旋耕和耙耕条件下为马唐、旱稗和牛筋草;常规耕作条件下,优势杂草为马唐、苘麻、旱稗和香附子。无秸秆还田条件下,免耕和常规耕作增加了杂草优势种的数量。秸秆全量还田后,免耕、耙耕和深松等耕作措施导致杂草群落的物种丰富度及均匀度均较高。无论哪种耕作条件,5a连续秸秆还田能够显著提高夏玉米籽粒产量和生物学产量,其中尤以常规耕作秸秆全量还田处理产量最高,且田间杂草的生物学产量与夏玉米的生物学产量呈显著负相关关系。     

Water use efficiency of a maize/cowpea intercrop on a highly acidic tropical soil as affected by liming and fertilizer application

Due to global warming, there is a need to increase the water use efficiency of crops under rainfed agriculture, particularly in semi-arid regions. Therefore, the effect of NPK fertilizer application (with or without liming) on the water use efficiency of a maize/cowpea intercropping system was investigated in the semi-arid part of Brazil. The crops were grown on a strongly acidic, sandy soil with three treatments: (i) Complete NPK fertilizer application with lime (Compl), (ii) Complete NPK fertilizer application without lime (Compl-L) and (iii) Control. On the average, dry matter production was 2.6 times higher with the Compl treatment than in the Control and 1.6 times higher than in the Compl-L treatment. The soil water balance was calculated with two different model approaches (HILLFLOW and EPICSEAR). When checked against measured soil water content during the growing period, both models produced accurate results, but only EPICSEAR was sensitive to the effects of liming and fertilizer application on soil water balance and dry matter production at this site. Comparison between the Compl and the Compl-L treatments shows that the increase in transpirational water use efficiency (WUE_T) (+63 and +80%, respectively) is mainly due to the application of NPK. Although the site is highly acid, liming was of minor importance for increasing the WUE_T. However, observations and simulations demonstrate that, through the additional application of lime, the gross water use efficiency (WUE_C) in a maize/cowpea intercropping system can be increased by 60% compared to sole application of NPK and by more than 160% compared to the control. Abbreviations: EPIC – erosion productivity impact calculator; EPICSEAR – erosion productivity impact calculator for semi-arid regions; TDR – time domain reflectometry; WUE – water use efficiency.     

Genome size and numerical polymorphism for the B chromosome in races of maize (Zea mays ssp. mays, Poaceae)

Twenty-one native populations (1120 individuals) of maize from Northern Argentina were studied. These populations, which belong to 13 native races, were cultivated at different altitudes (80-3620 m). Nineteen of the populations analyzed showed B chromosome (Bs) numerical polymorphism. The frequency of individuals with Bs varied from 0 to 94%. The number of Bs per plant varied from 0 to 8 Bs, with the predominant doses being 0, 1, 2, and 3. Those populations with varying number of Bs showed a positive and statistically significant correlation of mean number of Bs with altitude. The DNA content, in plants without Bs (A-DNA)(2n = 20), of 17 populations of the 21 studied was determined. A 36% variation (5.0-6.8 pg) in A-DNA content was found. A significant negative correlation between A-DNA content and altitude of cultivation and between A-DNA content and mean number of Bs was found. This indicates that there is a close interrelationship between the DNA content of A chromosomes and doses of Bs. These results suggest that there is a maximum limit to the mass of nuclear DNA so that Bs are tolerated as long as this maximum limit is not exceeded.     

Silicon promotes nodule formation and nodule function in symbiotic cowpea (Vigna unguiculata)

Applying silicon in the form of metasilicic acid (H₄SiO₃) or silicic acid (H₄SiO₃) to Bradyrhizobium -infected, hydroponically grown cowpea seedlings resulted in a significant ( P 0.05) increase in the number of nodules, nodule dry matter, and nitrogen fixed on a per plant basis. Total dry matter of plants increased with silicon supply, and the differences were significant ( P 0.05) at the higher silicon concentrations. Cowpea plants cultured in sand were also assessed for their response to silicic acid. Nodule number and nodule mass increased with silicon supply to sand cultured plants, though nitrogen fixation was unaltered. Although silicon is not essential for growth of cowpea, it is important for nodule formation and nodule functioning in hydroponically grown plants. Consequently, data collected and conclusions drawn from earlier glasshouse experiments, which have excluded silicon from nutrient solutions, may be flawed. Future studies on nodulation and nitrogen fixation using legumes in liquid culture must therefore include silicon as a nutrient element.     

Chlorophyll-deficient gene and morphological variations in Korean populations of maize (Zea mays)

We studied cultivated and naturalized Korean maize populations to determine the extent to which the chlorophylldeficient mutation and the phenotypic variations of two morphological characters (i.e., red coleoptiles and epicotyls, and the number of the first root hairs) are maintained. The frequency of the chlorophyll-deficient mutant gene (2.73% on average) was highly variable. Frequencies of red coleoptiles and epicotyls also were higher than expected from a mutation-selection balance. The average number of hairy phenotypes within populations was 1.8, ranging from 0.0 to 4.0. Naturalized populations were closely related to with cultivated communities. Most striking, however, was the more significant difference among populations than within populations with regard to both the frequency of chlorophyll-deficient mutant genes and the phenotypic variations of our two morphological characters. On a per-gene basis, the majority of the phenotypic variation (mean of 73.3%) resided among populations.     

Localization of the genes controlling B chromosome transmission rate in maize (Zea mays ssp. mays, Poaceae)

In previous papers we found that the frequency of B chromosomes in native races of maize varies considerably in different populations. Moreover, we found genotypes that control high and low transmission rates (TR) of B chromosomes in the Pisingallo race. In the present work crosses were made to determine whether the genes controlling B-TR are located on the normal chromosome set (As) or on the B chromosomes (Bs). We made female f.0B × male m.2B crosses between and within high (H) and low (L) B-TR groups. The Bs were transmitted on the male side in all cases. The mean B-TR from the progeny of f.0B (H) × m.2B (H) and f.0B (H) × m.2B (L) crosses was significantly higher than that from f.0B (L) × m.2B (L) and f.0B (L) × m.2B (H) crosses. The results show that the B-TR of the crosses corresponds to the H or L B-TR of the 0B female parents irrespective of the Bs of the male parent. This indicates that B-TR is genetically controlled by the 0B female parent and that these genes are located on the A chromosomes.     

弱光胁迫影响夏玉米光合效率的生理机制初探

大田条件下, 以普通夏玉米(Zea mays) ‘泰玉2号’为材料, 于授粉后1-20天遮光55% (+S), 以大田自然光照条件下生长的玉米作为对照(-S), 研究了遮光及恢复过程中玉米植株的光合性能、叶绿体荧光参数、叶黄素循环以及光能分配的变化, 初步揭示夏玉米开花后弱光条件下光适应的生理机制, 为玉米高产稳产提供理论依据。结果表明, 遮光后玉米穗位叶叶绿素含量及可溶性蛋白含量均减少, RuBP羧化酶和PEP羧化酶活性显著降低, 导致穗位叶净光合速率(P_n)迅速下降, 光饱和点也明显降低; 恢复初期P_n迅速升高, 光合关键酶活性有所增强。遮光后植株的最大光化学效率(F_v/F_m)、实际光化学效率(Ф_PSII)显著降低, 非光化学淬灭(NPQ)则显著升高, 而恢复初期植株穗位叶Ф_PSII有所升高, 表明突然暴露在自然光下的光合电子传递速率明显加快, 这与其光合速率及光合酶活性的趋势保持一致; 遮光处理对穗位叶叶黄素循环库的大小(紫黄质+花药黄质+玉米黄质(V + A + Z))影响不显著, 但使叶黄素循环的脱环氧化状态(A + Z)/(V + A + Z)增加; 遮光后植株分配于光化学反应的光能明显减少, 天线耗散光能比率显著增加, 恢复过程中植株主要以过剩非光化学反应的形式耗散过剩的光能。遮光后及恢复初期, 玉米植株的PSII原初光化学活性明显下降, 限制了光合碳代谢的电子供应从而抑制了光合作用, 主要依赖叶黄素循环途径进行能量耗散, 而在光照转换后遮光的玉米叶片在适应自然光过程中的光保护机制不断完善, 光合能力逐渐得到 恢复。     

Biochemical characterization of maize (Zea mays L.) for salt tolerance

The inherent differences for salt tolerance in two maize cultivars (Agatti-2002 and Sahiwal-2002) were evaluated in pot experiments. Plants were grown in half-strength of Hoagland nutrient solution added with 0, 80, 100, 120, 140 and 160 mM of NaCl. Salt stress markedly reduced the shoot and root lengths and fresh and dry masses. Reduction in growth attributes was more pronounced in cv. Agatti-2002 than cv. Sahiwal-2002. Both maize cultivars exhibited significant perturbations in important biochemical attributes being employed for screening the crops for salt tolerance. Cultivar Sahiwal-2002 was found salt tolerant as compared to cv. Agatti-2002 because it exhibited lower levels of H₂O₂, malondialdehyde (MDA) and higher activities of antioxidant enzymes. In addition, cultivar Sahiwal-2002 exhibited less salt-induced degradation of photosynthetic pigments, lower levels of toxic Na⁺ and Cl^?  and higher endogenous levels of K⁺ and K⁺/Na⁺ ratio. The results indicate that salt stress induced a marked increase in MDA, H₂O₂, relative membrane permeability, total soluble proteins and activities of antioxidant enzymes (superoxide dismutase, peroxidase, catalase andascorbate peroxidase). Moreover, increase in endogenous levels of Na⁺ and Cl^?  and decrease in K⁺ and K⁺/Na⁺ ratio and photosynthetic pigments were recorded in plants grown under salinity regimes.     

Genetics of resistance to the pink stem borer (Sesamia nonagrioides) in maize (Zea mays)

Our knowledge of the genetics of resistance to the pink stem borer ( Sesamia nonagrioides ) in maize ( Zea mays ) is restricted to a few crosses among maize inbreds. The objectives of this study were to enlarge our understanding of the genetics of traits related to damage by pink stem borer and yield under infestation and to use generation means analyses to compare per se and testcross performance for detecting epistatic effects. All generations, either per se or crossed to testers, were evaluated in a 10 × 10 triple lattice design under artificial infestation with S. nonagrioides in 2005 and 2006. Most traits fit an additive–dominance model; but evidence for epistasis for resistance and yield under infestation was shown. Epistasis, in general, did not appear to play an important role in the inheritance of yield under pink stem borer infestation. However, the epistasis contribution to maize yield performance could be important in some outstanding crosses such as EP42 × A637. Testcross generation means revealed epistatic effects undetected by the generation means analysis, but neither method was able to eliminate dominance effects that could prevail over epistatic effects.     

Chemical speciation of Zn in acidic soils: suitable soil extractant for assessing Zn availability to maize (Zea mays L.)

Abstract

Chemical fractions of soil Zn namely: water soluble (WS), exchangeable (EX), Pb displaceable (Pb-disp.), acid soluble (AS), Mn oxide occluded (MnOX), organically bound (OB), amorphous Fe oxide occluded (AFeOX), crystalline Fe oxide occluded (CFeOX), residual (RES) were determined in 20 surface (0–15 cm) samples of acidic soils from the provinces of Uttarakhand and Uttar Pradesh, India. The chemical fractions of soil Zn in acidic soils were found to be in the following descending order of Zn concentration: RES > CFeOX > Pb-Disp. > AFeOX > MnOX > AS > OB > EX > WS. These soil samples were also extracted by: DTPA (pH 7.3), DTPA (pH 5.3), AB-DTPA (pH 7.6), Mehlich 3 (pH 2.0), Modified Olsen, 0.01 N CaCl₂, 1 M MgCl ₂ and ion exchange resins. Chemical fractions and the soil extractable content of Zn estimated by different soil extractants were significantly correlated with some general soil properties. Maize (cv. Pragati) plants were grown in these soils for 35 days after emergence and Zn uptake by plants was compared with the amount of Zn extracted by different soil extractants and chemical fractions of Zn. Among chemical fractions of soil Zn, Pb-displaceable and acid soluble chemical fractions of soil Zn showed a significant and positive correlation with Zn uptake by maize. Path coefficient analysis also revealed that the acid soluble Zn fraction showed the highest positive and direct effect on Zn uptake (P=0.960). Among different multinutrient soil extractants evaluated for their suitability to assess Zn availability in acidic soils, DTPA (pH=5.3) was most suitable soil extractant, as the quantity of soil Zn extracted by this extractant showed a significant and positive correlation with the dry matter yield, Zn concentration and uptake by maize plants.     

Tribology of the root cap in maize (Zea mays) and peas (Pisum sativum)

Frictional resistance to a penetrating body can account for more than 80% of the total resistance to penetration of soil. We measured the frictional resistance between growing root caps of maize and pea and ground and smooth glass surfaces, which was linearly correlated to load, allowing calculation of the coefficient of kinetic friction and adhesion. Coefficients of kinetic friction between the root caps and the ground and smooth glass surfaces were approximately 0.04 and 0.02, respectively, the first measurements of the frictional properties of root tips at rates approaching those of root elongation, and an order of magnitude smaller than those previously reported. Results suggest that roots are well designed for penetrating soil, and encounter only small frictional resistance on the root cap. These data provide important parameters for modelling soil stresses and deformation around growing root tips.     

冬玉米对氮肥的吸收利用和需求

在不同施氮量下 ,研究了冬玉米对氮肥的吸收利用 ,结果表明 :(1 )冬玉米地上各部分中氮的累积随着用氮量的增加而增加 ,花丝期前的吸收量均多于后期 ,但高氮区的前期吸氮比大于低氮区 ;(2 )营养体氮的转移率随施氮量的增加而降低 ,但绝对量依然是高氮处理大于低氮处理 ,其中雄穗的转移率最高 ,叶的转移量最大 ;(3 )氮肥利用率随施氮量的增加而提高 ,但氮的生产力下降。根据试验结果 ,在肥力好的土地上种植冬玉米以 1 80～ 2 70 kg N/hm2比较适宜     

Root cortical aerenchyma inhibits radial nutrient transport in maize (Zea mays)

Background and Aims

Formation of root cortical aerenchyma (RCA) can be induced by nutrient deficiency. In species adapted to aerobic soil conditions, this response is adaptive by reducing root maintenance requirements, thereby permitting greater soil exploration. One trade-off of RCA formation may be reduced radial transport of nutrients due to reduction in living cortical tissue. To test this hypothesis, radial nutrient transport in intact roots of maize (Zea mays) was investigated in two radiolabelling experiments employing genotypes with contrasting RCA.

Methods

In the first experiment, time-course dynamics of phosphate loading into the xylem were measured from excised nodal roots that varied in RCA formation. In the second experiment, uptake of phosphate, calcium and sulphate was measured in seminal roots of intact young plants in which variation in RCA was induced by treatments altering ethylene action or genetic differences.

Key Results

In each of three paired genotype comparisons, the rate of phosphate exudation of high-RCA genotypes was significantly less than that of low-RCA genotypes. In the second experiment, radial nutrient transport of phosphate and calcium was negatively correlated with the extent of RCA for some genotypes.

Conclusions

The results support the hypothesis that RCA can reduce radial transport of some nutrients in some genotypes, which could be an important trade-off of this trait.     

Integration of Glomus mosseae with Chromolaena odorata powder for suppression of Meloidogyne incognita on maize (Zea mays L.)

Meloidogyne incognita infestation on maize results in heavy yield loss in farmers’ field. Most of the varieties adopted by subsistence farmers in Nigeria are susceptible to M. incognita. Beside these, the cost of control exceeding the profit from the crop using nematicides and the pollution risk they pose to the environment has necessitated the need for alternatives. Pot and field experiments were, therefore, conducted to investigate the effects of Chromolaena odorata powder and Glomus mosseae (a mycorrhizal fungus) on M. incognita pathogenicity on maize. Hybrid Oba super II improved maize variety adopted by local farmers was selected for the study. Maize plants were grown with G. mosseae (5 spores/g of soil) and soil amended with C. odorata powder (1% w/w) singly and in combination. Two weeks after emergence, Test plants were inoculated with 5000 M. incognita eggs. Sixty days after planting, destructive samples were assessed for root gall symptom and severity, and nematode population. Results show that G. mosseae and C. odorata powder were effective in controlling the population of M. incognita and the root knot nematode symptom and gave the highest yield parameters in combination. Single application of G. mosseae and C. odorata powder was similar in the effect on M. incognita and maize yield. Combination of G. mosseae and C. odorata powder may become a viable alternative to nematicide in managing M. incognita pathogenicity on maize as C. odorata powder may serve as a carrier medium for G. mosseae.