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近年来, 随着林下经济的蓬勃发展, 林下食用菌栽培业取得了较大的发展, 但栽培种类仅局限于已能进行大规模人工栽培的种类, 如香菇、平菇、杏鲍菇、木耳等。目前, 不能进行人工栽培却能真正与树木形成互惠互利关系的共生菌根食用菌栽培模式已成为研究方向, 这种栽培模式是“营建新型生态经济林——食用菌根菌经济林模式”[1], 即在营造经济林之前选择好树种和与其能共生的珍贵食用菌根菌种类, 人工使其合成菌根, 从单一生产林果为主转变为以生产林果和珍贵食用菌根菌为主, 既增加了经济收入, 又充分发挥了菌根共生关系的优势[2]。国际上主要通过培养菌根化幼苗, 建立种植园的方式在多种珍贵食用菌根菌生产上取得了成功, 而我国研究人员在黑孢块菌、乳牛肝菌、红汁乳菇等珍贵食用菌根菌栽培方面同样取得了较大进展[3-4]。目前, 食用菌根菌已逐渐成为农林业的新宠儿[5], 不仅因为这些食用菌根菌具有独特的食用和药用价值[6-7], 而且还能促进树木生长、提高产量、增强抗性、改良土壤。板栗(Castanea mollissima Bl.)是我国优良的经济林树种, 且能与多种珍贵食用菌根菌共生, 探索和发展板栗食用菌根菌生态林, 既在一定程度上解决我国某些地区板栗发展趋于饱和、缺乏经济增长的现状, 又能结合老龄板栗树的更新, 引入食用菌根菌栽培, 增加新的经济增长点。
发展板栗食用菌根菌经济林模式的基础是目的菌种能否与板栗人工合成菌根, 笔者在研制成功板栗食用菌根菌固体菌剂的基础上[8], 进行了板栗接种食用菌根菌菌根化效果试验, 以验证供试菌种能否侵染板栗根系, 能否形成菌根的典型结构——菌丝套和哈蒂氏网, 验证固体菌剂的有效性, 同时观测接种食用菌根菌对板栗幼苗生长的影响。
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育苗试验地位于浙江省诸暨市浬浦镇兼溪村(29°N, 120°E), 属亚热带季风气候区, 四季分明, 雨水较多, 光照充足。原为浙江省板栗主产区之一, 但近年来由于效益下降、林龄老化等原因, 板栗发展已停滞。本试验的最终目的就是引入食用菌根菌栽培, 以增加板栗种植效益。
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本文供试食用菌根菌及来源见表 1。菌种由中国林科院菌根研究开发中心经过固体发酵后制成固体菌剂, 用于育苗试验。
表 1 供试菌种名称及来源
Table 1. The name and sources of the tested strains
菌种名称Name of mycorrhiza fungi 菌株编号Strain number 菌种来源Strain sources 大红菇
(Russula alutacea Fr.)2014-10 中国林科院菌根研究开发中心采集、分离
Collection and separation of the research and development center of mycorrhizae, CAF.红绒盖牛肝菌
(Xerocomus chrysenteron Bull.ex Fr.)2014-14 中国林科院菌根研究开发中心采集、分离
Collection and separation of the research and development center of mycorrhizae, CAF.美味牛肝菌
(Boletus edulis Bull.Fr.)50559 中国农业菌种保藏中心提供
Provided by ACCC. -
测量和方差分析的结果(表 2、3)表明: 板栗幼苗苗高、地径各区组间差异不显著, 各处理间差异显著(P < 0.05); 板栗幼苗根系干质量区组间差异不显著, 各处理间差异极显著(P < 0.01)。
表 2 板栗幼苗苗高、地径、根系干质量方差分析结果
Table 2. Variation analysis of seedling height, basal diameter and root dry biomass of Chinese chestnut seedlings
变异来源
Sources of variation自由度
Freedom均方
Mean squareF F0.05 F0.01 苗高
Seedling height地径
Ground diameter根系干质量
Root dry weight区组Block 2 50.50 3.38 0.033 0.72 4.76 9.78 处理Processing 3 121.06 8.10* 6.10* 21.63** 误差Chance error 6 14.95 注: *表示差异显著(P < 0.05), **表示差异极显著(P < 0.01), 下同。
Note: * the difference was significant (P < 0.05), * * the difference was very significant (P < 0.01), the same below.表 3 板栗幼苗苗高、地径、根系干质量平均值及多重比较结果
Table 3. The multiple comparison of the average seedling height, basal diameter and root dry biomass of Chinese chestnut seedling
处理Processing 苗高Seedling height 地径Ground diameter 根系干质量Root dry weight 平均值/cm
Average/cm较CK提高/%
Improve than the CK/%平均值/mm
Average/mm较CK提高/%
Improve than the CK/%平均值/g
Average/g较CK提高/%
Improve than the CK/%大红菇Russula alutacea Fr. 60.8a 32.5 8.61a 19.4 19.97A 18.9 红绒盖牛肝菌Xerocomus chrysenteron Bull.ex Fr. 56.9a 24.0 8.23a 14.1 21.04A 25.3 美味牛肝菌Boletus edulis Bull.Fr. 56.3a 22.7 7.57b 5.0 19.16A 14.1 对照CK 45.9b - 7.21b - 16.79B - 注: 同列中不同小写字母表示差异显著, 不同大写字母表示差异极显著, 下同。
Note: in the same column, the different small letters indicate significant difference, the difference capital letters indicate very significant, the same below.由表 3可见: 各菌种处理苗高均显著高于对照, 大红菇、红绒盖牛肝菌和美味牛肝菌处理的平均苗高比对照分别提高32.5%、24.0%和22.7%, 各菌种间平均苗高差异不显著, 说明接种食用菌根菌菌种后, 对当年生幼苗的高生长有显著的促进作用; 大红菇和红绒盖牛肝菌处理幼苗的平均地径比对照分别提高19.4%和14.1%, 与对照差异显著, 而美味牛肝菌处理后幼苗的平均地径仅比对照提高5.0%, 二者间差异不显著。
大红菇、红绒盖牛肝菌和美味牛肝菌处理后根系平均干质量比对照分别提高18.9%、25.3%和14.1%, 均与对照差异极显著。
由表 4、5可见: 各菌种处理的侧根数和侧根总长均与对照差异极显著, 大红菇、红绒盖牛肝菌和美味牛肝菌处理平均侧根数分别比对照提高16.0%、14.1%和12.1%, 平均侧根总长分别比对照提高26.8%、28.9%和44.8%, 说明接种食用菌根菌对板栗幼苗根系的生长发育具有极显著的促进作用。由图 1(左为菌根化苗, 右为对照苗)可见: 大红菇处理的菌根化苗苗高、地径均优于对照, 根系比对照苗更发达, 侧根尤其是吸收根更多。图 2为菌种大红菇(2014-10)与板栗幼苗形成的菌根化根系, 图中箭头所指为吸收根所形成的白色菌根。
表 4 板栗幼苗侧根数及侧根总长方差分析结果
Table 4. Variation analysis of lateral roots number and lateral roots length of Chinese chestnut seedlings
变异来源
Sources of variation自由度
FreedomF F0.05 F0.01 侧根数
The number of lateral roots侧根总长
Lateral root length处理Processing 3 20.55** 27.6** 3.24 5.29 误差Chance error 16 表 5 板栗幼苗侧根数、侧根总长平均值及多重比较结果
Table 5. The multiple comparison of the average lateral roots number and lateral roots length of Chinese chestnut seedlings
处理Processing 侧根数The number of lateral roots 侧根总长Lateral root length 平均值/条
Average较CK提高/%
Improve than the CK/%平均值/cm
Average/cm较CK提高/%
Improve than the CK/%大红菇Russula alutacea Fr. 47.8A 16.0 410.0B 26.8 红绒盖牛肝菌Xerocomus chrysenteron Bull.ex Fr. 47.0A 14.1 416.6B 28.9 美味牛肝菌Boletus edulis Bull.Fr. 46.2A 12.1 468.2A 44.8 对照CK 41.2B - 323.3C - 图 1 2014-10处理菌根化幼苗与对照的比较
Figure 1. Comparison of mycorrhizal seedlings in the treatment of 2014-10 and control
图 2 2014-10与板栗形成的菌根化根系(箭头所指为菌根,下同)
Figure 2. 2014-10 and chestnut formation of mycorrhiza roots(The arrow refers to the mycorrhizal, the same below)
由图 3~5(左为菌根化根系, 右为对照根系)发现: 各菌种处理形成的菌根化根系不仅侧根多, 而且吸收根丰富, 并形成了大量的白色菌根。
图 3 菌种2014-10处理根系与对照的比较
Figure 3. Comparison of the root system in the treatment of 2014 - 10 and control.
图 4 菌种2014-14处理根系与对照的比较
Figure 4. Comparison of the root system in the treatment of 2014-14 and control.
图 5 菌种50559处理根系与对照的比较
Figure 5. Comparison of the root system in the treatment of 50559 and control.
综合以上调查结果, 生长指标在3个食用菌根菌间差异不显著, 说明这3个菌种对板栗幼苗生长指标的促进作用一致。
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通过对各处理幼苗根系的观察发现: 3个菌种均与板栗根系形成了菌根。利用目测法和计数统计法统计幼苗菌根化率和吸收根菌根化率。由表 6可见: 接种食用菌根菌后, 各菌种均与板栗幼苗根系形成菌根, 幼苗平均菌根化率均高于80%, 吸收根平均菌根化率均≥50%。由图 6~8可见: 大红菇(2014-10)、红绒盖牛肝菌(2014-14)和美味牛肝菌(50559)与板栗形成菌根化根系, 大量的吸收根形成了白色的菌根。3个菌种处理板栗幼苗的部分吸收根特写照片(图 9~11)表明: 吸收根基本形成了白色菌根, 说明板栗幼苗根系的菌根化程度较高[2]。对照根系也有少量的菌根, 这可能是土壤中的土著菌根菌造成的, 由于其菌的数量较少, 因此, 菌根化率均显著低于人工接种, 虽然幼苗菌根化率达14.8%, 但平均吸收根菌根化率只有3%, 基本可以忽略不计。
表 6 板栗幼苗菌根化率和吸收根菌根化率调查结果
Table 6. The investigation of the average mycorrhization rate of seedlings and absorbing roots of Chinese chestnut seedlings
处理
Processing幼苗平均菌根化率/%
Average infection rate of seedlings/%吸收根平均菌根化率/%
Average infection rate of absorbing root大红菇Russula alutacea Fr. 95.8 57 红绒盖牛肝菌
Xerocomus chrysenteron Bull.ex Fr.87.0 53 美味牛肝菌
Boletus edulis Bull.Fr.96.7 50 CK 14.8 3 -
选取典型菌根, 通过肉眼或体视镜观察发现, 3个菌种与板栗幼苗根系形成的菌根均可见明显的菌丝套结构, 外延菌丝较多(表 7)。选取典型菌根, 制作石蜡切片, 经显微镜观察, 吸收根横切片及纵切片上均可见菌丝束缠绕在吸收根尖表面, 即形成了菌丝套。同时在根皮层外侧细胞间隙亦可见明显的菌丝侵入, 从菌根的横切片上可观察到念珠状结构, 即形成了典型的哈蒂氏网结构。由图 12~14可以清晰地观察到根表皮外包裹的菌丝套和根皮层外侧细胞间的哈蒂氏网结构。可见, 3种食用菌根菌菌种均与板栗幼苗根系共生并形成了典型的菌根结构——菌丝套和哈蒂氏网, 表明板栗与3种食用菌根菌人工合成菌根成功。详细的微观结构观察将另文描述。
表 7 不同菌种处理的板栗幼苗根系形成的菌根外观形态
Table 7. The morphological characteristics of mycorhiza of Chinese chestnut seedlings inoculated with different solid inocula
菌种
Strain菌根颜色
Mycorrhizal color外观描述
Mycorrhizal appearance大红菇
Russula alutacea Fr.白色
White顶端稍膨大, 以二叉状、塔状为主, 具有明显的菌丝套, 外延菌丝多。
There is swelling on the top of the mycorrhizae, with more two fork and tower shaped, with the clearly mycelial mantle, and more external mycelium.红绒盖牛肝菌
Xerocomus chrysenteron Bull.ex Fr白色
White顶端基本无膨大, 以二叉状和单棒状为主, 具有明显的菌丝套, 外延菌丝较多。
There is no swelling on the top of the mycorrhizae, with two fork and single rod like, with the clearly mycelial mantle, and more external mycelium.美味牛肝菌
Boletus edulis Bull.Fr.白色
White顶端基本无膨大, 以二叉状、羽状为主, 具有明显的菌丝套, 外延菌丝较多。
There is no swelling on the top of the mycorrhizae, with two fork and feather dominated like, with the clearly mycelial mantle, and more external mycelium.图 12 2014-10形成的菌根根尖横切片显微照片(×400)
Figure 12. Microscopic photographs of the apical transverse sections of the mycorrhiza root formed with strain 2014-10(×400)
板栗接种食用菌根真菌的菌根化效果及对幼苗促生效应的研究
Study of Mycorrization of Inoculating Edible Mycorrhizal Fungi on Chinese Chestnut and Its Effect on Promoting Seedling Growth
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摘要:
目的 通过板栗幼苗人工接种食用菌根菌固体菌剂,分析其对板栗幼苗生长指标和根系菌根化的效果,以验证大红菇(菌株编号2014-10)、红绒盖牛肝菌(菌株编号2014-14)和美味牛肝菌(菌株编号50559)这3种食用菌根菌固体菌剂作为板栗接种体的有效性及其促生效应,为进一步探索“板栗食用菌根菌新型经济林栽培模式”提供基础。 方法 在板栗播种时分别接种上述3种食用菌根菌固体菌剂,并设空白作对照。对当年生幼苗苗高、地径、根系干质量、侧根数、侧根总长、幼苗侵染率、吸收根侵染率等指标进行调查和分析,并制作菌根石蜡切片进行微观结构观察。 结果 表明:(1)上述3种食用菌根菌在人工接种下均能侵染板栗幼苗根系,平均幼苗菌根化率分别达95.8%、87.0%和96.7%,平均吸收根菌根化率分别达57%、53%和50%,达较高的侵染强度;经微观观察,板栗幼苗根系均形成典型的菌根结构——菌丝套和哈蒂氏网。(2)与对照幼苗相比,3个菌种处理幼苗的平均苗高分别提高32.5%、24.0%和22.7%,平均地径分别提高19.4%、14.1%和5.0%,差异显著(P < 0.05);平均根系干质量分别提高18.9%、25.3%和14.1%,平均侧根数分别提高16.0%、14.1%和12.1%,平均侧根总长分别提高26.8%、28.9%和44.8%,差异极显著(P < 0.01)。 结论 (1) 上述供试食用菌根菌均与板栗根系建立了共生关系,幼苗菌根化效果良好,吸收根菌根化强度高,形成了菌根的典型结构,即菌丝套和哈蒂氏网,验证了上述菌种固体菌剂的有效性,认为其可以作为板栗食用菌根菌栽培的接种体而用于生产实践,为探索“板栗食用菌根菌新型经济林栽培模式”提供了基础。(2)上述供试菌种接种后,对板栗幼苗各项主要生长指标均有不同程度的促进作用,说明上述食用菌根菌对板栗幼苗具有显著的促生效应。 Abstract:Objective The objective of this study is to verify the effectiveness of three edible mycorrhizal fungi species (Russula alutacea, Xerocomus chrysenteron, and Boletus edulis) as solid inocula and their effect on promoting the growth of Chinese chestnut (Castanea mollissima) seedlings, so as to provide some references for the study of "new non-timber forest cultivation model of Chinese chestnut- edible mycorrhizal fungi". Method The three species of mycorrhizal fungi were inoculated when sowing Chinese chestnut. The seedling height, basal diameter, root dry biomass, lateral root number, lateral root length, seedling infection rate, infection rate of absorption root were investigated and analyzed. Result The results show that: (1) The 3 mycorrhizal fungi species could infect the root of Chinese chestnut seedling under artificial inoculation condition, the average infection rates of the 3 mycorrhizal fungi species were 95.8%, 87.0% and 96.7% respectively, the average infection rates of absorption root were 57%, 53% and 50% respectively, showing that the infection rate reached a high level. The results of microscopic observation showed that the typical structure of the mycelial mantle and Hartig net were formed. (2) Compared with the control seedlings, the average heights of treated seedlings increased by 32.5%, 23.9% and 22.7% respectively, the average diameters increased by 19.4%, 14.1% and 5.0% respectively. significantly differences were observed between treated seedlings and control (P < 0.05). The average root dry biomass increased by 18.9%, 25.3% and 14.1% respectively, the average number of lateral roots increased by 16%, 14.1% and 12.1% respectively, the average length of lateral roots increased by 26.8%, 28.9% and 44.8% respectively, highly significantly differences were observed between treated seedlings and control (P < 0.01). Conclusion (1) The 3 mycorrhizal fungi species established a symbiotic relation with Chinese chestnut root, and formed a typical structure of mycorrhizae, i.e. the mycelial mantle and Hartig net. The results verified the effectiveness of the solid edible mycorrhizal inocula. It was proved that they can be used as inocula for the Chinese chestnut cultivation. It provided the references for the study of the "new non-timber forest cultivation model of Chinese chestnut-edible mycorrhizal fungi". (2) In the study, the main growth indexes of Chinese chestnut seedlings were promoted in various degrees, which showed that the edible mycorrhizal fungi had a significant effect on promoting the growth of Chinese chestnut seedlings. -
Key words:
- Castanea mollissima
- / edible mycorrhizal fungi
- / mycorrhization
- / growth effects
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表 1 供试菌种名称及来源
Table 1. The name and sources of the tested strains
菌种名称Name of mycorrhiza fungi 菌株编号Strain number 菌种来源Strain sources 大红菇
(Russula alutacea Fr.)2014-10 中国林科院菌根研究开发中心采集、分离
Collection and separation of the research and development center of mycorrhizae, CAF.红绒盖牛肝菌
(Xerocomus chrysenteron Bull.ex Fr.)2014-14 中国林科院菌根研究开发中心采集、分离
Collection and separation of the research and development center of mycorrhizae, CAF.美味牛肝菌
(Boletus edulis Bull.Fr.)50559 中国农业菌种保藏中心提供
Provided by ACCC.表 2 板栗幼苗苗高、地径、根系干质量方差分析结果
Table 2. Variation analysis of seedling height, basal diameter and root dry biomass of Chinese chestnut seedlings
变异来源
Sources of variation自由度
Freedom均方
Mean squareF F0.05 F0.01 苗高
Seedling height地径
Ground diameter根系干质量
Root dry weight区组Block 2 50.50 3.38 0.033 0.72 4.76 9.78 处理Processing 3 121.06 8.10* 6.10* 21.63** 误差Chance error 6 14.95 注: *表示差异显著(P < 0.05), **表示差异极显著(P < 0.01), 下同。
Note: * the difference was significant (P < 0.05), * * the difference was very significant (P < 0.01), the same below.表 3 板栗幼苗苗高、地径、根系干质量平均值及多重比较结果
Table 3. The multiple comparison of the average seedling height, basal diameter and root dry biomass of Chinese chestnut seedling
处理Processing 苗高Seedling height 地径Ground diameter 根系干质量Root dry weight 平均值/cm
Average/cm较CK提高/%
Improve than the CK/%平均值/mm
Average/mm较CK提高/%
Improve than the CK/%平均值/g
Average/g较CK提高/%
Improve than the CK/%大红菇Russula alutacea Fr. 60.8a 32.5 8.61a 19.4 19.97A 18.9 红绒盖牛肝菌Xerocomus chrysenteron Bull.ex Fr. 56.9a 24.0 8.23a 14.1 21.04A 25.3 美味牛肝菌Boletus edulis Bull.Fr. 56.3a 22.7 7.57b 5.0 19.16A 14.1 对照CK 45.9b - 7.21b - 16.79B - 注: 同列中不同小写字母表示差异显著, 不同大写字母表示差异极显著, 下同。
Note: in the same column, the different small letters indicate significant difference, the difference capital letters indicate very significant, the same below.表 4 板栗幼苗侧根数及侧根总长方差分析结果
Table 4. Variation analysis of lateral roots number and lateral roots length of Chinese chestnut seedlings
变异来源
Sources of variation自由度
FreedomF F0.05 F0.01 侧根数
The number of lateral roots侧根总长
Lateral root length处理Processing 3 20.55** 27.6** 3.24 5.29 误差Chance error 16 表 5 板栗幼苗侧根数、侧根总长平均值及多重比较结果
Table 5. The multiple comparison of the average lateral roots number and lateral roots length of Chinese chestnut seedlings
处理Processing 侧根数The number of lateral roots 侧根总长Lateral root length 平均值/条
Average较CK提高/%
Improve than the CK/%平均值/cm
Average/cm较CK提高/%
Improve than the CK/%大红菇Russula alutacea Fr. 47.8A 16.0 410.0B 26.8 红绒盖牛肝菌Xerocomus chrysenteron Bull.ex Fr. 47.0A 14.1 416.6B 28.9 美味牛肝菌Boletus edulis Bull.Fr. 46.2A 12.1 468.2A 44.8 对照CK 41.2B - 323.3C - 表 6 板栗幼苗菌根化率和吸收根菌根化率调查结果
Table 6. The investigation of the average mycorrhization rate of seedlings and absorbing roots of Chinese chestnut seedlings
处理
Processing幼苗平均菌根化率/%
Average infection rate of seedlings/%吸收根平均菌根化率/%
Average infection rate of absorbing root大红菇Russula alutacea Fr. 95.8 57 红绒盖牛肝菌
Xerocomus chrysenteron Bull.ex Fr.87.0 53 美味牛肝菌
Boletus edulis Bull.Fr.96.7 50 CK 14.8 3 表 7 不同菌种处理的板栗幼苗根系形成的菌根外观形态
Table 7. The morphological characteristics of mycorhiza of Chinese chestnut seedlings inoculated with different solid inocula
菌种
Strain菌根颜色
Mycorrhizal color外观描述
Mycorrhizal appearance大红菇
Russula alutacea Fr.白色
White顶端稍膨大, 以二叉状、塔状为主, 具有明显的菌丝套, 外延菌丝多。
There is swelling on the top of the mycorrhizae, with more two fork and tower shaped, with the clearly mycelial mantle, and more external mycelium.红绒盖牛肝菌
Xerocomus chrysenteron Bull.ex Fr白色
White顶端基本无膨大, 以二叉状和单棒状为主, 具有明显的菌丝套, 外延菌丝较多。
There is no swelling on the top of the mycorrhizae, with two fork and single rod like, with the clearly mycelial mantle, and more external mycelium.美味牛肝菌
Boletus edulis Bull.Fr.白色
White顶端基本无膨大, 以二叉状、羽状为主, 具有明显的菌丝套, 外延菌丝较多。
There is no swelling on the top of the mycorrhizae, with two fork and feather dominated like, with the clearly mycelial mantle, and more external mycelium. -
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