外生菌根真菌氮、钾营养特性及其对汞胁迫的反应
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摘要
氮和钾均是树木需要量最多的必需元素,对森林的发生、发展、演替和衰亡具有极其重要的作用。在森林土壤中,绝大多数氮是植物不能直接吸收的有机氮,有效氮通常处于缺乏状态。我国南方有大面积的红壤、黄壤和石灰岩土壤,由于受到强烈淋溶等多种因素的影响,土壤有效钾含量一般较低,大部分钾是难于利用的无效钾。森林一般不施肥,主要依靠土壤供应植物的氮、钾营养。此外,汞污染是目前全球性的普遍问题,全世界每年因人类活动而释放的汞超过2909吨,中国是最大的汞污染源。在汞污染严重的地区,抑制树根吸收氮、钾等养分,妨碍树木生长发育和天然更新,最终导致森林退化和衰亡。菌根是植物根系与菌根真菌形成的共生体,是植物与真菌长期共同进化的结果。优良的外生菌根真菌与木本植物的根系形成外生菌根后,能促进树木的生长,改善它们的营养状况,提高其抗逆(干旱、病害、重金属等)能力。所以,研究外生菌根真菌的氮、钾营养特性及其对汞胁迫的反应,对于我国天然林的保护和人工林的营造,以及汞污染的土地上植树造林具有重要意义。
试验利用源于我国南北方和国外的14株外生菌根真菌为材料。供试菌株为：彩色豆马勃Pt715(采自四川西昌桉树林)[Pisolithus tinctorius (Pers.) Coker & Couch (Strain Pt 715)],松乳菇Ld-1、Ld-2和Ld-3(采自重庆缙云山马尾松林)[Lactarius deliciosus (L.:Fr.) Gray (strain Ld-1, Ld-2 & Ld-3)],硬皮马勃Sp01(采自重庆金佛山马尾松林)[Scleroderma polyrhizum Pers. (strain Sp-01)],牛肝菌Bo 02(采自重庆金佛山马尾松林)[Boletus spp. (strain Bo-02)],褐环乳牛肝菌Sl07和Sl14(采自重庆金佛山马尾松林)[Suillus luteus(L.:Fr.) Gray (strain Sl 07 & Sl 14)],黄空柄牛肝菌Gc 99(采自内蒙古大青山)[Gyroporus cyanescens(Bull.:Fr.) Quel.(strain Gc 99)],土生空团菌Cg04(采自内蒙古大青山)[Cenococcum geophilum Fr. (strain Cg 04)],厚环乳牛肝菌Sg03(采自内蒙古大青山)[Suillus grevillei (Kl.)Sing. (strain Sg 03)],网纹灰包Lp01(采自内蒙古大青山)[Lycoperdon perlatum Pers (strain Sp 01)],乳牛肝菌Sb05(采自内蒙古大青山)[Suillus bovinus (L.:Fr.)Kuntze (strain Sb 05)],以及土生空团菌Cg SIV(采自西班牙欧洲火炬松林)[Cenococcum geophilum Fr. (Strain Cg SIV)]等。在Pachlewski液体培养基中,提供不同的氮源,加入不同浓度的钾,以及设置不同程度的汞胁迫,研究了外生菌根真菌对不同氮源(氨态氮、硝态氮、尿素和蛋白质)的利用能力；不同供钾条件下,外生菌根真菌的生长、氮磷钾吸收、有机酸分泌,草酸代谢关键酶的活性；以及汞对外生菌根真菌生长、氮磷钾吸收、有机酸分泌、氮素利用酶活性等的影响。获得主要结论如下：
1.1多数外生菌根真菌在以铵态氮为氮源的培养基中生长良好,蛋白质氮其次,尿素和硝态氮对生长的促进作用相对较差。说明供试菌株对不同的氮源既具有偏嗜性,同时也具有多样性,可适应多种不同的生态条件。在植树造林工作中,应充分考虑土壤—菌株适配性。在供试菌株体内,均能检测到蛋白酶、几丁质酶、脲酶和硝酸还原酶,不同菌株之间酶活性差异显著。在外生菌根真菌体内,氮素利用酶的存在与活性高低是它们利用不同氮源的生理基础。
1.2培养基中的钾离子浓度显著影响外生菌根真菌的生长。无钾生长不良,多数情况下中钾浓度适合外生菌根真菌生长。在供应低钾时,Sp01和Pt 715的生长量最大,中钾和高钾时降低,表明Sp01和Pt715可在低钾条件下正常生长,可能比较适宜缺钾的土壤。钾对外生菌根真菌含氮量的影响因菌株而异,一般以中钾或高钾时含氮量较高。除Sl14以外,培养基中较高的钾浓度在一定程度上会抑制外生菌根真菌对磷的吸收。培养基钾浓度是决定外生菌根真菌含钾量的直接原因,供钾水平的升高促进外生菌根真菌吸收钾。
1.3 K+显著影响外生菌根真菌分泌草酸、乙酸和H+。无钾或低钾时草酸和乙酸均分泌最多,中钾或高钾时减少；多数菌株无钾或低钾时H+分泌最多,但Bo 02和Lp01、Pt715和Sb05分别在高钾和中钾时H+分泌量最大。外生菌根真菌能分泌较多的草酸、乙酸和H+可能有利于含钾矿物的风化,促进矿物结构钾和层间钾的释放,可视为外生菌根真菌活化土壤无效钾的机理之一。
1.4缺钾提高外生菌根真菌苹果酸脱氢酶、乙醛酸脱氢酶和草酰乙酸酶的活性,其变化趋势与类似草酸分泌。推测在供钾较低时,促进外生菌根真菌草酸合成关键酶基因的表达。此外,不同菌株苹果酸脱氢酶、乙醛酸脱氢酶和草酰乙酸酶活性差异较大,推测在不同菌株体内中,上述草酸代谢途径酶的相对重要性有所不同。
1.5在0～150gmol Hg·L-1的范围内,汞对Cg SIV生长的抑制作用不明显,在高汞浓度的培养基中生物量仅比对照减少9.7%,外生菌根真菌具有较高的耐汞性；汞虽然使Pt 715和Ld-3的生物量有所降低,但它们的生物量仍然较高。在汞胁迫条件下,外生菌根真菌的含氮量有升有降,磷、钾含量总体上增加。氮、磷、钾的吸收量Pt715和Ld-3最多。
1.6在汞胁迫条件下,蛋白酶、几丁质酶、脲酶和硝酸还原酶的活性不同,推测外生菌根真菌可能合成对汞敏感性不同的等位酶。此外,5μmol Hg2+.L-1～50μmol Hg2+.L-1的培养基中,有关氮素利用酶的活性无显著变化甚至有所提高,推测在一定的汞污染条件下,对外生菌根真菌氮素利用能力无显著影响。值得注意的是,在正常和汞胁迫条件下,Pt 715和Ld-3的蛋白酶、脲酶、硝酸还原酶和几丁质酶的活性均最高,表现出较强的氮素利用能力。推断在汞污染的土壤上种植桉树和松树,接种Pt 715和Ld-3可能改善寄主植物的氮素营养。
1.7在汞胁迫条件下,外生菌根真菌分泌草酸具有普遍性,菌株Sp01、Sl14和Cg04能分泌酒石酸,Sp01、Cg04和Pt715能分泌丁二酸。草酸、酒石酸和琥珀酸分泌量均表现出高汞>中汞>低汞>无汞的规律。由于草酸、酒石酸和琥珀酸具有络合汞的能力,降低外界环境和菌丝体内活性汞的浓度,故在汞胁迫的条件下外生菌根真菌分泌有机酸可视为抗(耐)汞危害的机理之一。
Nitrogen and potassium, both of which play critical roles in origin, development, succession and decline of forests, are most needed essential elements by trees. In forest ecosystems, the exchangeable N in the soil solution is very low relative to total soil N and most of the soil nitrogen is in organic form. Large areas of red, yellow and limestone soil that distribute in the south of China are leaching strongly, thus the readily available potassium in soil is also very low. Since fertilization is difficult in forest, nitrogen and potassium nutrition of plants are mainly supplied by soil. On the other hand, mercury pollution becomes a global problem due to anthropic mercury release, which is more than 2909 tons per year with the largest part contributed by China. In areas with severe mercury pollution, absorption of nutrients including nitrogen and potassium by plants is restrained, development and renewal of trees are disturbed, and ultimately lead to forest degradation and decline. Ectomycorrhizal fungi are symbiotically associated with the roots of many woody plants and can dramatically enhance plant growth. Ectomycorrhiza-forming fungi alter the nutrient-acquisition capabilities of vascular plants, and may play an important role in tolerance of plant to drought, disease and heavy metal. Therefor, studying nitrogen and potassium nutrition of ectomycorrhizal fungi and their response to mercury stress is important to natural forest protection, afforestation and tree planting on mercury contaminated soil.
14 ectomycorrhizal fungal strains involved in this study, including Pisolithus tinctorius (Pers.) Coker & Couch (Strain Pt 715, isolated from soils under eucalyptus, Xichang, Sichuan province), Lactarius deliciosus (L.:Fr.) Gray (strain Ld-1, Ld-2 & Ld-3, isolated from soils under masson pine, Jinyun mountain, Chongqin), Scleroderma polyrhizum Pers. (strain Sp-01, isolated from soils under masson pine, Jinfo mountain, Chongqin), Scleroderma polyrhizum Pers. (strain Sp-01, isolated from soils under masson pine, Jinfo mountain, Chongqin), Boletus spp. (strain Bo-02, isolated from soils under masson pine, Jinfo mountain, Chongqin), Suillus luteus(L.:Fr.) Gray (strain Sl 07 & Sl 14, isolated from soils under masson pine, Jinfo mountain, Chongqin), Gyroporus cyanescens(Bull.:Fr.) Quel.(strain Gc 99, isolated from Daqing mountain, Inner Mongolia), Cenococcum geophilum Fr. (strain Cg 04, isolated from Daqing mountain, Inner Mongolia), Suillus grevillei (Kl.)Sing. (strain Sg 03, isolated from Daqing mountain, Inner Mongolia), Lycoperdon perlatum Pers (strain Lp 01, isolated from Daqing mountain, Inner Mongolia), Suillus bovinus (L.:Fr.)Kuntze (strain Sb 05, isolated from Daqing mountain, Inner Mongolia), Cenococcum geophilum Fr. (Strain Cg SIV, from Spain). Ectomycorrhizal fungi were cultured in Pachlewski liquid media with different nitrogen resources, different potassium concentrations or different mercury concentrations. Nitrogen utilization to different nitrogen resources (NH_4~+-N, NO_3~--N, urea and protein)by ectomycorrhizal fungi, growth rate, absorption of nitrogen, potassium and phosphorus, organic acids secretion and activities of key enzymes in oxalate metabolism pathways of ectomycorrhizal fungi under different potassium concentrations, as well as effects of mercury on growth rate, absorption of nitrogen, phosphorus and potassium, organic acids secretion and activities of nitrogen utilizing enzymes of ectomycorrhizal fungi, were studied. The result showed:
Growth rate of most ectomycorrhizal fungi strains was promoted by ammonium as nitrogen resource. Protein was in the next place, whereas urea and nitrate performed poorly in promoting growth of ectomycorrhizal fungi. It is implied that the tested strains might not only benefit to nitrogen utilization diversity of host plants, but also favored different nitrogen sources. It suggested that the adaptation between soil and ectomycorrhizal fungal strains should be considered during afforestation. All of the four tested strains expressed protease, chitinase, urease and nitrate reductase, and there were significant differences among them. The existence and activity of nitrogen utilizing enzymes in the mycelia are the physiological basis of utilizing of different nitrogen resources by ectomycorrhizal fungi.
Potassium concentration in the media influenced the growth rate of ectomycorrhizal fungi significantly. Growth was restrained without potassium, while most strains growed optimized in the concentration of middle-potassium. Sp 01 and Pt 715 got the fastest growth rate in low-potassium media, which suggested that Sp 01 and Pt 715 might both adapt to soils short of potassium. Effects of potassium on the nitrogen concentration of ectomycorrhizal fungi differed between strains. Higher concentration of potassium restrained phosphorus absorption by ectomycorrhizal fungi to some extents except SI 14. potassium concenteation in the media determined the potassium content in ectomycorrhizal fungal hyphae, as the increase of potassium level promoted potassium absorption by ectomycorrhizal fungi.
K~+ affected secretion of oxalate, acetic acid and H~+ significantly. The secretion of oxalate and acetic acid reached the highest in no-K~+ or low-K~+ media, while was the lowest in middle or high-K~+ media. Most strains secreted H~+ at the utmost in no-K~+ or low-K~+ media, but Bo 02 and Lp 01, Pt 715 and Sb 05 secreted H~+ at the utmost in middle or high-K~+ media, respectively. Oxalate, acetic acid and proton secreted by ectomycorrhizal fungi may benefit to mineral weathering and the release of K trapped between clay layers.
Activities of malate dehydrogenase, glyoxylate dehydrogenase and oxaloacetate hydrase in ectomycorrhizal fungi increased when lack of K, which showed some similarities with oxalate secretion. It suggested that low concentration of potassium may promote the expression of key enzymes in oxlate metabolism pathways. Furthermore, Activities of malate dehydrogenase, glyoxylate dehydrogenase and oxaloacetate hydrase in different strains differed siglificantly, which may imply the relative importance of these oxalate metabolism enzymes is different in different strains.
Growth of fungal hyphae was restrained by mercury, whereas Cg SIV showed tolerance to Hg in some extent, with the least decrease on biomass. Although the biomass of Pt 715 and Ld-3 decreased by mercury treatment, their biomass are still the highest among tested strains.
Effects of Hg on activities of nitrogen utilizing enzymes in ectomycorrhizal fungi varied with strains, enzyme categories and Hg~(2+) concentration. Since the inherited characteristics differ among strains, enzyme synthesis may vary under condition of the mercury existence, and express allozymes with different sensitivity to Hg. Moreover, low to medium Hg concentration promoted or did not affect the activities of nitrogen utilizing enzymes, which suggested that nitrogen utilization by ectomycorrhizal fungi might not be restraint or even be promoted. Activities of protease, urease, nitrate reductase and chitinase were the highest in Pt15 and Ld-3 with or without Hg treatment, which showed relatively high capacity of nitrogen utilization. Inoculating Pt715 and Ld-3 may improve nitrogen nutrition of eucalyptuses and pines planting in mercury contaminated soils.
Under mercury stress, all of the tested strains secreted oxalate, while Sp 01, Sl 14 and Cg 04 secreted tartaric acid, and Sp 01, Cg 04 and Pt 715 secreted succinic acid. The efflux rate of oxalate, tartaric acid and succinic acid all showed the order of high Hg> middle Hg> low Hg> no Hg. Oxalate, tartaric acid and succinic acid are chelators of heavy metals, thus may reduce mercury concentration in the hyphae and the environment.
试验利用源于我国南北方和国外的14株外生菌根真菌为材料。供试菌株为：彩色豆马勃Pt715(采自四川西昌桉树林)[Pisolithus tinctorius (Pers.) Coker & Couch (Strain Pt 715)],松乳菇Ld-1、Ld-2和Ld-3(采自重庆缙云山马尾松林)[Lactarius deliciosus (L.:Fr.) Gray (strain Ld-1, Ld-2 & Ld-3)],硬皮马勃Sp01(采自重庆金佛山马尾松林)[Scleroderma polyrhizum Pers. (strain Sp-01)],牛肝菌Bo 02(采自重庆金佛山马尾松林)[Boletus spp. (strain Bo-02)],褐环乳牛肝菌Sl07和Sl14(采自重庆金佛山马尾松林)[Suillus luteus(L.:Fr.) Gray (strain Sl 07 & Sl 14)],黄空柄牛肝菌Gc 99(采自内蒙古大青山)[Gyroporus cyanescens(Bull.:Fr.) Quel.(strain Gc 99)],土生空团菌Cg04(采自内蒙古大青山)[Cenococcum geophilum Fr. (strain Cg 04)],厚环乳牛肝菌Sg03(采自内蒙古大青山)[Suillus grevillei (Kl.)Sing. (strain Sg 03)],网纹灰包Lp01(采自内蒙古大青山)[Lycoperdon perlatum Pers (strain Sp 01)],乳牛肝菌Sb05(采自内蒙古大青山)[Suillus bovinus (L.:Fr.)Kuntze (strain Sb 05)],以及土生空团菌Cg SIV(采自西班牙欧洲火炬松林)[Cenococcum geophilum Fr. (Strain Cg SIV)]等。在Pachlewski液体培养基中,提供不同的氮源,加入不同浓度的钾,以及设置不同程度的汞胁迫,研究了外生菌根真菌对不同氮源(氨态氮、硝态氮、尿素和蛋白质)的利用能力；不同供钾条件下,外生菌根真菌的生长、氮磷钾吸收、有机酸分泌,草酸代谢关键酶的活性；以及汞对外生菌根真菌生长、氮磷钾吸收、有机酸分泌、氮素利用酶活性等的影响。获得主要结论如下：
1.1多数外生菌根真菌在以铵态氮为氮源的培养基中生长良好,蛋白质氮其次,尿素和硝态氮对生长的促进作用相对较差。说明供试菌株对不同的氮源既具有偏嗜性,同时也具有多样性,可适应多种不同的生态条件。在植树造林工作中,应充分考虑土壤—菌株适配性。在供试菌株体内,均能检测到蛋白酶、几丁质酶、脲酶和硝酸还原酶,不同菌株之间酶活性差异显著。在外生菌根真菌体内,氮素利用酶的存在与活性高低是它们利用不同氮源的生理基础。
1.2培养基中的钾离子浓度显著影响外生菌根真菌的生长。无钾生长不良,多数情况下中钾浓度适合外生菌根真菌生长。在供应低钾时,Sp01和Pt 715的生长量最大,中钾和高钾时降低,表明Sp01和Pt715可在低钾条件下正常生长,可能比较适宜缺钾的土壤。钾对外生菌根真菌含氮量的影响因菌株而异,一般以中钾或高钾时含氮量较高。除Sl14以外,培养基中较高的钾浓度在一定程度上会抑制外生菌根真菌对磷的吸收。培养基钾浓度是决定外生菌根真菌含钾量的直接原因,供钾水平的升高促进外生菌根真菌吸收钾。
1.3 K+显著影响外生菌根真菌分泌草酸、乙酸和H+。无钾或低钾时草酸和乙酸均分泌最多,中钾或高钾时减少；多数菌株无钾或低钾时H+分泌最多,但Bo 02和Lp01、Pt715和Sb05分别在高钾和中钾时H+分泌量最大。外生菌根真菌能分泌较多的草酸、乙酸和H+可能有利于含钾矿物的风化,促进矿物结构钾和层间钾的释放,可视为外生菌根真菌活化土壤无效钾的机理之一。
1.4缺钾提高外生菌根真菌苹果酸脱氢酶、乙醛酸脱氢酶和草酰乙酸酶的活性,其变化趋势与类似草酸分泌。推测在供钾较低时,促进外生菌根真菌草酸合成关键酶基因的表达。此外,不同菌株苹果酸脱氢酶、乙醛酸脱氢酶和草酰乙酸酶活性差异较大,推测在不同菌株体内中,上述草酸代谢途径酶的相对重要性有所不同。
1.5在0～150gmol Hg·L-1的范围内,汞对Cg SIV生长的抑制作用不明显,在高汞浓度的培养基中生物量仅比对照减少9.7%,外生菌根真菌具有较高的耐汞性；汞虽然使Pt 715和Ld-3的生物量有所降低,但它们的生物量仍然较高。在汞胁迫条件下,外生菌根真菌的含氮量有升有降,磷、钾含量总体上增加。氮、磷、钾的吸收量Pt715和Ld-3最多。
1.6在汞胁迫条件下,蛋白酶、几丁质酶、脲酶和硝酸还原酶的活性不同,推测外生菌根真菌可能合成对汞敏感性不同的等位酶。此外,5μmol Hg2+.L-1～50μmol Hg2+.L-1的培养基中,有关氮素利用酶的活性无显著变化甚至有所提高,推测在一定的汞污染条件下,对外生菌根真菌氮素利用能力无显著影响。值得注意的是,在正常和汞胁迫条件下,Pt 715和Ld-3的蛋白酶、脲酶、硝酸还原酶和几丁质酶的活性均最高,表现出较强的氮素利用能力。推断在汞污染的土壤上种植桉树和松树,接种Pt 715和Ld-3可能改善寄主植物的氮素营养。
1.7在汞胁迫条件下,外生菌根真菌分泌草酸具有普遍性,菌株Sp01、Sl14和Cg04能分泌酒石酸,Sp01、Cg04和Pt715能分泌丁二酸。草酸、酒石酸和琥珀酸分泌量均表现出高汞>中汞>低汞>无汞的规律。由于草酸、酒石酸和琥珀酸具有络合汞的能力,降低外界环境和菌丝体内活性汞的浓度,故在汞胁迫的条件下外生菌根真菌分泌有机酸可视为抗(耐)汞危害的机理之一。
Nitrogen and potassium, both of which play critical roles in origin, development, succession and decline of forests, are most needed essential elements by trees. In forest ecosystems, the exchangeable N in the soil solution is very low relative to total soil N and most of the soil nitrogen is in organic form. Large areas of red, yellow and limestone soil that distribute in the south of China are leaching strongly, thus the readily available potassium in soil is also very low. Since fertilization is difficult in forest, nitrogen and potassium nutrition of plants are mainly supplied by soil. On the other hand, mercury pollution becomes a global problem due to anthropic mercury release, which is more than 2909 tons per year with the largest part contributed by China. In areas with severe mercury pollution, absorption of nutrients including nitrogen and potassium by plants is restrained, development and renewal of trees are disturbed, and ultimately lead to forest degradation and decline. Ectomycorrhizal fungi are symbiotically associated with the roots of many woody plants and can dramatically enhance plant growth. Ectomycorrhiza-forming fungi alter the nutrient-acquisition capabilities of vascular plants, and may play an important role in tolerance of plant to drought, disease and heavy metal. Therefor, studying nitrogen and potassium nutrition of ectomycorrhizal fungi and their response to mercury stress is important to natural forest protection, afforestation and tree planting on mercury contaminated soil.
14 ectomycorrhizal fungal strains involved in this study, including Pisolithus tinctorius (Pers.) Coker & Couch (Strain Pt 715, isolated from soils under eucalyptus, Xichang, Sichuan province), Lactarius deliciosus (L.:Fr.) Gray (strain Ld-1, Ld-2 & Ld-3, isolated from soils under masson pine, Jinyun mountain, Chongqin), Scleroderma polyrhizum Pers. (strain Sp-01, isolated from soils under masson pine, Jinfo mountain, Chongqin), Scleroderma polyrhizum Pers. (strain Sp-01, isolated from soils under masson pine, Jinfo mountain, Chongqin), Boletus spp. (strain Bo-02, isolated from soils under masson pine, Jinfo mountain, Chongqin), Suillus luteus(L.:Fr.) Gray (strain Sl 07 & Sl 14, isolated from soils under masson pine, Jinfo mountain, Chongqin), Gyroporus cyanescens(Bull.:Fr.) Quel.(strain Gc 99, isolated from Daqing mountain, Inner Mongolia), Cenococcum geophilum Fr. (strain Cg 04, isolated from Daqing mountain, Inner Mongolia), Suillus grevillei (Kl.)Sing. (strain Sg 03, isolated from Daqing mountain, Inner Mongolia), Lycoperdon perlatum Pers (strain Lp 01, isolated from Daqing mountain, Inner Mongolia), Suillus bovinus (L.:Fr.)Kuntze (strain Sb 05, isolated from Daqing mountain, Inner Mongolia), Cenococcum geophilum Fr. (Strain Cg SIV, from Spain). Ectomycorrhizal fungi were cultured in Pachlewski liquid media with different nitrogen resources, different potassium concentrations or different mercury concentrations. Nitrogen utilization to different nitrogen resources (NH_4~+-N, NO_3~--N, urea and protein)by ectomycorrhizal fungi, growth rate, absorption of nitrogen, potassium and phosphorus, organic acids secretion and activities of key enzymes in oxalate metabolism pathways of ectomycorrhizal fungi under different potassium concentrations, as well as effects of mercury on growth rate, absorption of nitrogen, phosphorus and potassium, organic acids secretion and activities of nitrogen utilizing enzymes of ectomycorrhizal fungi, were studied. The result showed:
Growth rate of most ectomycorrhizal fungi strains was promoted by ammonium as nitrogen resource. Protein was in the next place, whereas urea and nitrate performed poorly in promoting growth of ectomycorrhizal fungi. It is implied that the tested strains might not only benefit to nitrogen utilization diversity of host plants, but also favored different nitrogen sources. It suggested that the adaptation between soil and ectomycorrhizal fungal strains should be considered during afforestation. All of the four tested strains expressed protease, chitinase, urease and nitrate reductase, and there were significant differences among them. The existence and activity of nitrogen utilizing enzymes in the mycelia are the physiological basis of utilizing of different nitrogen resources by ectomycorrhizal fungi.
Potassium concentration in the media influenced the growth rate of ectomycorrhizal fungi significantly. Growth was restrained without potassium, while most strains growed optimized in the concentration of middle-potassium. Sp 01 and Pt 715 got the fastest growth rate in low-potassium media, which suggested that Sp 01 and Pt 715 might both adapt to soils short of potassium. Effects of potassium on the nitrogen concentration of ectomycorrhizal fungi differed between strains. Higher concentration of potassium restrained phosphorus absorption by ectomycorrhizal fungi to some extents except SI 14. potassium concenteation in the media determined the potassium content in ectomycorrhizal fungal hyphae, as the increase of potassium level promoted potassium absorption by ectomycorrhizal fungi.
K~+ affected secretion of oxalate, acetic acid and H~+ significantly. The secretion of oxalate and acetic acid reached the highest in no-K~+ or low-K~+ media, while was the lowest in middle or high-K~+ media. Most strains secreted H~+ at the utmost in no-K~+ or low-K~+ media, but Bo 02 and Lp 01, Pt 715 and Sb 05 secreted H~+ at the utmost in middle or high-K~+ media, respectively. Oxalate, acetic acid and proton secreted by ectomycorrhizal fungi may benefit to mineral weathering and the release of K trapped between clay layers.
Activities of malate dehydrogenase, glyoxylate dehydrogenase and oxaloacetate hydrase in ectomycorrhizal fungi increased when lack of K, which showed some similarities with oxalate secretion. It suggested that low concentration of potassium may promote the expression of key enzymes in oxlate metabolism pathways. Furthermore, Activities of malate dehydrogenase, glyoxylate dehydrogenase and oxaloacetate hydrase in different strains differed siglificantly, which may imply the relative importance of these oxalate metabolism enzymes is different in different strains.
Growth of fungal hyphae was restrained by mercury, whereas Cg SIV showed tolerance to Hg in some extent, with the least decrease on biomass. Although the biomass of Pt 715 and Ld-3 decreased by mercury treatment, their biomass are still the highest among tested strains.
Effects of Hg on activities of nitrogen utilizing enzymes in ectomycorrhizal fungi varied with strains, enzyme categories and Hg~(2+) concentration. Since the inherited characteristics differ among strains, enzyme synthesis may vary under condition of the mercury existence, and express allozymes with different sensitivity to Hg. Moreover, low to medium Hg concentration promoted or did not affect the activities of nitrogen utilizing enzymes, which suggested that nitrogen utilization by ectomycorrhizal fungi might not be restraint or even be promoted. Activities of protease, urease, nitrate reductase and chitinase were the highest in Pt15 and Ld-3 with or without Hg treatment, which showed relatively high capacity of nitrogen utilization. Inoculating Pt715 and Ld-3 may improve nitrogen nutrition of eucalyptuses and pines planting in mercury contaminated soils.
Under mercury stress, all of the tested strains secreted oxalate, while Sp 01, Sl 14 and Cg 04 secreted tartaric acid, and Sp 01, Cg 04 and Pt 715 secreted succinic acid. The efflux rate of oxalate, tartaric acid and succinic acid all showed the order of high Hg> middle Hg> low Hg> no Hg. Oxalate, tartaric acid and succinic acid are chelators of heavy metals, thus may reduce mercury concentration in the hyphae and the environment.
引文
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