土沉香（Aquilaria sinensis （Lour.）Gilg）结香机制的研究
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摘要
土沉香(Aquilaria sinensis (Lour.) Gilg)为我国特有的生产高品质香料及国药沉香的植物，关于沉香的形成过程，国内学者以土沉香为研究对象做了一些相关研究，国外不少学者则以沉香属其它的种为研究对象，如Aquilaria agallocha、Aquilaria malaccensis、Aquilaria crassna等，也做了大量的相关研究。本研究从土沉香与真菌的关系入手，从植物生理的角度，开展了真菌的分离鉴定与分析、真菌对土沉香木材组织成分及化学性质的影响、土沉香与结香真菌互作中的生理代谢变化等方面的研究，试图解决沉香结香过程中的几个关键问题。主要的研究结果如下：
1.土沉香(Aquilaria sinensis (Lour.) Gilg)树干受到创伤之后，包括可可毛壳色单隔孢(Lasiodiplodia theobromae)、茄病镰刀菌(Fusarium solani)、弯孢菌(Curvularia spp.)和拟盘多孢(Pestalotiopsis microspora)等致病菌种类会随之增加。这些致病菌在未创伤树干组织中并未发现，说明这些致病菌是通过创伤伤口侵入，物理创伤能够结香是多种致病菌侵染共同作用的结果。
2.在不同结香程度样品中，真菌种类不尽相同，且按未结香→始结香→已结香顺利依次减少。说明沉香物质的形成对有些真菌有抑制作用，使得一些真菌难以生存，其它的真菌取而代之，沉香的形成过程并不是某些真菌自始至终的结果。在有沉香物质的部分，即始结香部分和已结香部分，都发现了致病菌层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)。
3.土沉香(Aquilaria sinensis (Lour.) Gilg)树干滴注多变根毛霉(Rhizomucorvariabilis)、层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)三种菌剂后，脂肪酸的下降出现相似的规律，即前期（尤其是前2个月）下降较快，后期下降平稳，其中十八碳烯-[9]-酸下降幅度最大，其次是十八碳二烯-[10,2]-酸，棕榈酸降幅最小；三种菌剂中，多变根毛霉(Rhizomucor variabilis)对土沉香中脂肪酸成份的破坏最大，红褐肉座菌(Hypocrea jecorina)次之，而层出镰刀菌(Fusarium proliferatum)最小。
4.土沉香(Aquilaria sinensis (Lour.) Gilg)树干滴注多变根毛霉(Rhizomucorvariabilis)、层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)三种菌剂后，6个月时层出镰刀菌(Fusarium proliferatum)组苄基丙酮含量最高，达到104.7ug·g~(-1)，其次是红褐肉座菌(Hypocrea jecorina)组（33.38ug·g~(-1)），而多变根毛霉(Rhizomucorvariabilis)组极其微量，仅有1.94ug·g~(-1)。说明真菌种类对苄基丙醇含量的影响很大，多变根毛霉(Rhizomucor variabilis)对苄基丙酮的生成作用甚微，而层出镰刀菌(Fusariumproliferatum)效果最佳。
5.本研究通过对比液体菌剂和真菌培养液对苄基丙酮含量的影响，发现无菌的真菌培养液与液体菌剂间对苄基丙酮含量的影响并无显著差异。说明，沉香的形成与真菌并无直接关系，但与真菌代谢物关系密切。
6.土沉香(Aquilaria sinensis (Lour.) Gilg)树干滴注层出镰刀菌(Fusarium proliferatum)培养液6个月后，木材结香部分热水抽提物含量较白木部分增高了26.923%，说明层出镰刀菌培养液致使木材组织细胞壁受损，代谢物增多；木材结香部分苯-醇抽提物增高184.665%，为白木部分的2.85倍，表明木材中部分糖类化合物在木材向沉香变化过程中发生了降解作用，而萜类化合物、芳香族化合物等有机物含量随之增加；结香部分综纤维素含量比白木部分减少了21.737%，半纤维素减少9.986%，而木质素含量增加了38.316%，说明层出镰刀菌培养液可以降解综纤维素中的某些化学成分，但对木材的主要化学成分木质素无降解作用；与白木部分比较，结香部分木材组织中S元素增加了24.85%，其它矿物元素出现了不同程度的减少(7.92%~55.96%)，说明木材组织受到了胁迫；溶性糖的含量增高了56.10%，淀粉下降了62.92%，说明土沉香(Aquilaria sinensis (Lour.)Gilg)正常的生长与呼吸平衡受到了损坏，致使淀粉向糖转化。
7.沉香物质主要出现在导管和木射线薄壁组织。本研究的石蜡切片中沉香物质多的地方没有发现任何真菌。发现真菌菌丝体的地方沉香物质并不多，且所发现的菌不是研究中所采用的层出镰刀菌(Fusarium proliferatum)。说明，沉香物质不是层出镰刀菌(Fusarium proliferatum)在土沉香(Aquilaria sinensis (Lour.) Gilg)木材组织中生长繁殖的代谢产物。
8.添加层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)菌液后，两组土沉香(Aquilaria sinensis (Lour.) Gilg)组培幼苗叶片细胞膜脂氧化、细胞膜透性和糖代谢均存在明显差异。从细胞膜脂氧化来看，更有利于沉香物质形成的层出镰刀菌(Fusarium proliferatum)引发的胁迫反应中叶片细胞膜脂氧化（MDA）较红褐肉座菌(Hypocrea jecorina)轻，尤其在前期(24h～48h)；从细胞膜透性(相对电导率)的变化来看，层出镰刀菌组呈现逐渐上升的趋势，红褐肉座菌组则呈现先快后慢的上升趋势，说明层出镰刀菌(Fusarium proliferatum)对细胞膜的损害是一个渐近式的过程，而红褐肉座菌(Hypocrea jecorina)的损害更剧烈；从可溶性糖来看，两者在24h内均变化不大，48h时均出现了剧增，但红褐肉座菌组明显高于层出镰刀菌组，而72h后红褐肉座菌组下降幅度大于层出镰刀菌组，说明红褐肉座菌(Hypocrea jecorina)的侵染导致了糖代谢的不规律变化，而层出镰刀菌(Fusarium proliferatum)侵染后糖代谢相对平稳。
9.从活性氧代谢来看，滴注两个结香真菌菌液后，在短期(0~24h)内土沉香(Aquilariasinensis (Lour.) Gilg)组培幼苗叶片均出现了活性氧剧增的现象，之后均持续上升，但增幅存在差异，导致后期（96h时）经红褐肉座菌(Hypocrea jecorina)菌液处理的土沉香组培幼苗O_2~-和H_2O_2分别高于层出镰刀菌(Fusarium proliferatum)4.42U·g~(-1)和26.55U·g~(-1)。两个结香真菌对土沉香组培幼苗叶片中活性氧清除酶活性的影响亦存在差异，主要表现在两方面：一方面表现在活性上，层出镰刀菌(Fusarium proliferatum)处理后活性氧清除酶活性（SOD、POD和CAT）要高于红褐肉座菌(Hypocrea jecorina)；另一面表现在变化规律上，层出镰刀菌(Fusarium proliferatum)处理后活性氧清除酶活性的变化较缓，而红褐肉座菌(Hypocrea jecorina)的变化更为剧烈。
10.以层出镰刀菌培养液为母液，根据生理胁迫形成沉香的机理，添加NaCl（X1）和茉莉酸甲酯（X2），研究结香诱导剂配比。采用均匀设计及多元逐步回归分析，结果表明：茉莉酸甲酯对沉香醇溶性浸出物含量的影响没有NaCl大，但其在低水平条件下有利于提高沉香醇溶性浸出物含量；以母液+1.8248%NaCl+0.0046%茉莉酸甲酯为结香诱导剂进行滴注，1年后沉香中醇溶性浸出物（Y）获得理论最大值13.6012%。
Aquilaria sinensis (Lour.) Gilg was a paticular tree for producing agarwood used ashigh-quality incense and traditional chinese medicine in china. At abroad, there were somestudies on agar formating mechanism of Aquilaria agallocha, Aquilaria malaccensis, Aquilariacrassna, etc.. In china, studies targeted on Aquilaria sinensis (Lour.) Gilg have been carried outsince1970s. In this paper, studies on agar formation mechanism started with relation betweenAquilaria sinensis (Lour.) Gilg and the involved fungi from the standpoint of plant physiology,which including separation, purification and identification on involved fungi, fungi effect onchemical composition and chemical property of the wood tissue, physiological and metabolicchanges in the interaction between Aquilaria sinensis (Lour.) Gilg and the fungi. The mainresults about agar formation were as follows:
1. After wounding Aquilaria sinensis (Lour.) Gilg on the trunk, more pathogenic fungiwould appear, including Lasiodiplodia theobromae, Fusarium solani, Curvularia spp. andPestalotiopsis Microspora, etc.. The above pathogenic fungi were not found in the healthytrunk. It indicated that these pathogenic fungi were from outside through wound, and agarformation by physical wounding was the interaction results of various pathogenic fungiinfection.
2. In the studies, it was found that fungal species respectively decreased in the order：no-agar wood→little-agar wood→agar wood. It indicated that the formation of agar materialhad inhibiting effect on certain fungi, and then other fungi would replace them, so the agarformation process was not the result of some specific fungi infecting from first to last.Fusarium proliferatum and Hypocrea jecorina were found in the two test sample: little-agarwood, agar wood.
3. After instilling with the liquid inoculum of Rhizomucor variabilis, Fusariumproliferatum and Hypocrea jecorina, fatty acid in the timber of Aquilaria sinensis (Lour.) Gilgdecreased by similar rule, early (especially in the first2months) decreasing rapidly, and then steadily decreasing. The fatty acid consisted of octadecenic-[9]-acid, octadecenic-[10,2]-acidand palmitinic acid, among which, octadecenic-[9]-acid decreased at most, followed byoctadecenic-[10,2]-acid, and palmitic acid decreased at least. So the three fungi all coulddecompose the fatty acid in the timber of Aquilaria sinensis (Lour.) Gilg, the decompositiondegree in the order：Rhizomucor variabilis→Hypocrea jecorina→Fusarium proliferatum.
4. From the view of the content of benzyl acetone, on the6thmonth, benzyl acetonecontent of the sample instilled with liquid Fusarium proliferatum inoculum was the highest,reaching104.7μg·g~(-1), followed by the sample instilled with liquid Hypocrea jecorina inoculum(33.38μg·g~(-1)), and the sample instilled with liquid Hypocrea jecorina inoculum was very low,only1.94μg·g~(-1). It had been shown that the fungus species had great effect on benzyl alcoholcontent, namely, Rhizomucor variabilis had little effect on the formation of benzyl acetone, andFusarium proliferatum the best effect.
5. In this study, the effect of liquid inoculant on benzyl acetone content was comparedwith sterile fungal culture solution. The results had shown that there was little differencebetween liquid inoculant and sterile fungal culture solution. It indicated that there was notdirect relationship between agar formation and the fungal mycelium, on the contrary, there wasclose relationship between agar formation and mycelium metabolites.
6. After instilled with the culture solution of Fusarium proliferatum, on the6thmonthhot-water extractives content in the agar wood was higher than that in no-agar wood by26.923%, which indicated that the cell wall of the timber tissue had been partially damaged andthe metabolites increased; Benzene-ethanol extractives in the agar wood was higher than that inno-agar wood by184.665%, which showed that a part of carbohydrate had been degradated,terpenoid compounds, aromatic compounds content increased accordingly; Holocellulosecontent in the agar wood was lower than that in no-agar wood by21.737%, pentosan(hemicellulose) also lower by9.986%, and lignose content higher by38.316%, which indicatedthat the culture solution could degradate Holocellulose, but not degradate lignose; Comparedwith the no-agar wood, mineral elements content, except for S(increased by24.85%), in theagar wood decreased by7.92%~55.96%, which showed that the timber tissue was under stress, the S content increasing may be the result of passive absorption for self-protection; In theprocess of agar formation, normal balance between growth and respiration had been striked,which resulted in starch conversing to sugar, so the soluble sugar content in the agarwoodincreased by56.10%and starch content decreased by62.92%.
7. Agar material occurred mainly in parenchyma cell of the xylem tracheal and xylem rays.In paraffin sections, no fungi were found where abundant agar engendered, and agar was littlewhere the mycelium was found. Moreover, the fungus found in paraffin sections was notFusarium proliferatum. It had been shown that agar was not metabolites of growth andreproduction of Fusarium proliferatum in timber tissue of Aquilaria sinensis (Lour.) Gilg.
8. After Added Aquilaria sinensis (Lour.) Gilg tissue culture seedling with culture solutionof Fusarium proliferatum and Hypocrea jecorina, there was significant difference in cellmembrane lipid oxidation, cell membrane permeability and glucose metabolism between thetwo groups of samples. From the view of cell membrane lipid peroxidation, the MDA contentin leaves of the samples added Hypocrea jecorina culture solution (H-group) increasedsignificantly, while the one of the samples added Fusarium proliferatum culture solution(F-group) increased less, especially in early phase (24h~48h), which showed that Fusariumproliferatum, as a fungus more conducive to agar formation, could cause Aquilaria sinensis(Lour.) Gilg physiologic stress reaction, but the cell membrane lipid oxidation was not asserious as Hypocrea jecorina. From the view of cell membrane permeability changes, REC(Relative Electrical Conductivity) in the leaves of H-group increased steadily, while F-group sREC increased rapidly at first and then increases slowly. which showed that culture solution ofFusarium proliferatum would damage leaves s cell membrane gradually, while culturesolution of Hypocrea jecorina would damage acutely; From the view of soluble sugar, theleaves’ soluble-sugar in the two groups of samples all changed little in24h, and then allincreased rapidly with H-group more quickly, at the48thhour soluble-sugar content in theH-group samples was significantly higher than that in F-group samples, after the72thhour, thesoluble-sugar content in H-group samples declined more than that in F-group samples, whichindicated that Hypocrea jecorina infection caused nutrient transport disorder and resulted in irregular change of glucose metabolism, while Fusarium proliferatum effect on glucosemetabolism relatively stable.
9. After Added Aquilaria sinensis (Lour.) Gilg tissue culture seedling with culture solutionof Fusarium proliferatum and Hypocrea jecorina, ROS surge phenomenon all occurred in ashort period (0~24h). Later, at the96thhour the superoxide radical and hydrogen peroxide inthe H-group samples were higher than that of F-group samples by4.42U·g~(-1)and26.55U·g~(-1).There were significant differences in active-oxygen scavenging enzymes during thephysiologic stress phase caused by Fusarium proliferatum and Hypocrea jecorina. Thedifferences were manifested in two aspects：firstly, the enzymes activity (including SOD, PODand CAT) of F-group samples were higher than that of H-group samples；Secondly, changeregularity of the enzymes activity was different, enzymes activity in F-group samples changedrelative steadily, while the one in H-group changed acutely.
10. In this study, the hypothesis “physiological stress causing agar” was brought forward.Based on the hypothesis, a trial about agar revulsant formula, including Fusarium proliferatumculture solution, NaCl and JAs, was carried out according to uniform design. The results hadbeen shown that NaCl had greater effect on agarwood quanlity than JAs; the best agarrevulsant formula was Fusarium proliferatum culture solution added1.8248%NaCl and0.0046%JAs. Instilling the trunk with the agar revulsant formula, one year later, maximizedalcohol-soluble extractive content in the wood would be13.6012%theoretically.
1.土沉香(Aquilaria sinensis (Lour.) Gilg)树干受到创伤之后，包括可可毛壳色单隔孢(Lasiodiplodia theobromae)、茄病镰刀菌(Fusarium solani)、弯孢菌(Curvularia spp.)和拟盘多孢(Pestalotiopsis microspora)等致病菌种类会随之增加。这些致病菌在未创伤树干组织中并未发现，说明这些致病菌是通过创伤伤口侵入，物理创伤能够结香是多种致病菌侵染共同作用的结果。
2.在不同结香程度样品中，真菌种类不尽相同，且按未结香→始结香→已结香顺利依次减少。说明沉香物质的形成对有些真菌有抑制作用，使得一些真菌难以生存，其它的真菌取而代之，沉香的形成过程并不是某些真菌自始至终的结果。在有沉香物质的部分，即始结香部分和已结香部分，都发现了致病菌层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)。
3.土沉香(Aquilaria sinensis (Lour.) Gilg)树干滴注多变根毛霉(Rhizomucorvariabilis)、层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)三种菌剂后，脂肪酸的下降出现相似的规律，即前期（尤其是前2个月）下降较快，后期下降平稳，其中十八碳烯-[9]-酸下降幅度最大，其次是十八碳二烯-[10,2]-酸，棕榈酸降幅最小；三种菌剂中，多变根毛霉(Rhizomucor variabilis)对土沉香中脂肪酸成份的破坏最大，红褐肉座菌(Hypocrea jecorina)次之，而层出镰刀菌(Fusarium proliferatum)最小。
4.土沉香(Aquilaria sinensis (Lour.) Gilg)树干滴注多变根毛霉(Rhizomucorvariabilis)、层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)三种菌剂后，6个月时层出镰刀菌(Fusarium proliferatum)组苄基丙酮含量最高，达到104.7ug·g~(-1)，其次是红褐肉座菌(Hypocrea jecorina)组（33.38ug·g~(-1)），而多变根毛霉(Rhizomucorvariabilis)组极其微量，仅有1.94ug·g~(-1)。说明真菌种类对苄基丙醇含量的影响很大，多变根毛霉(Rhizomucor variabilis)对苄基丙酮的生成作用甚微，而层出镰刀菌(Fusariumproliferatum)效果最佳。
5.本研究通过对比液体菌剂和真菌培养液对苄基丙酮含量的影响，发现无菌的真菌培养液与液体菌剂间对苄基丙酮含量的影响并无显著差异。说明，沉香的形成与真菌并无直接关系，但与真菌代谢物关系密切。
6.土沉香(Aquilaria sinensis (Lour.) Gilg)树干滴注层出镰刀菌(Fusarium proliferatum)培养液6个月后，木材结香部分热水抽提物含量较白木部分增高了26.923%，说明层出镰刀菌培养液致使木材组织细胞壁受损，代谢物增多；木材结香部分苯-醇抽提物增高184.665%，为白木部分的2.85倍，表明木材中部分糖类化合物在木材向沉香变化过程中发生了降解作用，而萜类化合物、芳香族化合物等有机物含量随之增加；结香部分综纤维素含量比白木部分减少了21.737%，半纤维素减少9.986%，而木质素含量增加了38.316%，说明层出镰刀菌培养液可以降解综纤维素中的某些化学成分，但对木材的主要化学成分木质素无降解作用；与白木部分比较，结香部分木材组织中S元素增加了24.85%，其它矿物元素出现了不同程度的减少(7.92%~55.96%)，说明木材组织受到了胁迫；溶性糖的含量增高了56.10%，淀粉下降了62.92%，说明土沉香(Aquilaria sinensis (Lour.)Gilg)正常的生长与呼吸平衡受到了损坏，致使淀粉向糖转化。
7.沉香物质主要出现在导管和木射线薄壁组织。本研究的石蜡切片中沉香物质多的地方没有发现任何真菌。发现真菌菌丝体的地方沉香物质并不多，且所发现的菌不是研究中所采用的层出镰刀菌(Fusarium proliferatum)。说明，沉香物质不是层出镰刀菌(Fusarium proliferatum)在土沉香(Aquilaria sinensis (Lour.) Gilg)木材组织中生长繁殖的代谢产物。
8.添加层出镰刀菌(Fusarium proliferatum)和红褐肉座菌(Hypocrea jecorina)菌液后，两组土沉香(Aquilaria sinensis (Lour.) Gilg)组培幼苗叶片细胞膜脂氧化、细胞膜透性和糖代谢均存在明显差异。从细胞膜脂氧化来看，更有利于沉香物质形成的层出镰刀菌(Fusarium proliferatum)引发的胁迫反应中叶片细胞膜脂氧化（MDA）较红褐肉座菌(Hypocrea jecorina)轻，尤其在前期(24h～48h)；从细胞膜透性(相对电导率)的变化来看，层出镰刀菌组呈现逐渐上升的趋势，红褐肉座菌组则呈现先快后慢的上升趋势，说明层出镰刀菌(Fusarium proliferatum)对细胞膜的损害是一个渐近式的过程，而红褐肉座菌(Hypocrea jecorina)的损害更剧烈；从可溶性糖来看，两者在24h内均变化不大，48h时均出现了剧增，但红褐肉座菌组明显高于层出镰刀菌组，而72h后红褐肉座菌组下降幅度大于层出镰刀菌组，说明红褐肉座菌(Hypocrea jecorina)的侵染导致了糖代谢的不规律变化，而层出镰刀菌(Fusarium proliferatum)侵染后糖代谢相对平稳。
9.从活性氧代谢来看，滴注两个结香真菌菌液后，在短期(0~24h)内土沉香(Aquilariasinensis (Lour.) Gilg)组培幼苗叶片均出现了活性氧剧增的现象，之后均持续上升，但增幅存在差异，导致后期（96h时）经红褐肉座菌(Hypocrea jecorina)菌液处理的土沉香组培幼苗O_2~-和H_2O_2分别高于层出镰刀菌(Fusarium proliferatum)4.42U·g~(-1)和26.55U·g~(-1)。两个结香真菌对土沉香组培幼苗叶片中活性氧清除酶活性的影响亦存在差异，主要表现在两方面：一方面表现在活性上，层出镰刀菌(Fusarium proliferatum)处理后活性氧清除酶活性（SOD、POD和CAT）要高于红褐肉座菌(Hypocrea jecorina)；另一面表现在变化规律上，层出镰刀菌(Fusarium proliferatum)处理后活性氧清除酶活性的变化较缓，而红褐肉座菌(Hypocrea jecorina)的变化更为剧烈。
10.以层出镰刀菌培养液为母液，根据生理胁迫形成沉香的机理，添加NaCl（X1）和茉莉酸甲酯（X2），研究结香诱导剂配比。采用均匀设计及多元逐步回归分析，结果表明：茉莉酸甲酯对沉香醇溶性浸出物含量的影响没有NaCl大，但其在低水平条件下有利于提高沉香醇溶性浸出物含量；以母液+1.8248%NaCl+0.0046%茉莉酸甲酯为结香诱导剂进行滴注，1年后沉香中醇溶性浸出物（Y）获得理论最大值13.6012%。
Aquilaria sinensis (Lour.) Gilg was a paticular tree for producing agarwood used ashigh-quality incense and traditional chinese medicine in china. At abroad, there were somestudies on agar formating mechanism of Aquilaria agallocha, Aquilaria malaccensis, Aquilariacrassna, etc.. In china, studies targeted on Aquilaria sinensis (Lour.) Gilg have been carried outsince1970s. In this paper, studies on agar formation mechanism started with relation betweenAquilaria sinensis (Lour.) Gilg and the involved fungi from the standpoint of plant physiology,which including separation, purification and identification on involved fungi, fungi effect onchemical composition and chemical property of the wood tissue, physiological and metabolicchanges in the interaction between Aquilaria sinensis (Lour.) Gilg and the fungi. The mainresults about agar formation were as follows:
1. After wounding Aquilaria sinensis (Lour.) Gilg on the trunk, more pathogenic fungiwould appear, including Lasiodiplodia theobromae, Fusarium solani, Curvularia spp. andPestalotiopsis Microspora, etc.. The above pathogenic fungi were not found in the healthytrunk. It indicated that these pathogenic fungi were from outside through wound, and agarformation by physical wounding was the interaction results of various pathogenic fungiinfection.
2. In the studies, it was found that fungal species respectively decreased in the order：no-agar wood→little-agar wood→agar wood. It indicated that the formation of agar materialhad inhibiting effect on certain fungi, and then other fungi would replace them, so the agarformation process was not the result of some specific fungi infecting from first to last.Fusarium proliferatum and Hypocrea jecorina were found in the two test sample: little-agarwood, agar wood.
3. After instilling with the liquid inoculum of Rhizomucor variabilis, Fusariumproliferatum and Hypocrea jecorina, fatty acid in the timber of Aquilaria sinensis (Lour.) Gilgdecreased by similar rule, early (especially in the first2months) decreasing rapidly, and then steadily decreasing. The fatty acid consisted of octadecenic-[9]-acid, octadecenic-[10,2]-acidand palmitinic acid, among which, octadecenic-[9]-acid decreased at most, followed byoctadecenic-[10,2]-acid, and palmitic acid decreased at least. So the three fungi all coulddecompose the fatty acid in the timber of Aquilaria sinensis (Lour.) Gilg, the decompositiondegree in the order：Rhizomucor variabilis→Hypocrea jecorina→Fusarium proliferatum.
4. From the view of the content of benzyl acetone, on the6thmonth, benzyl acetonecontent of the sample instilled with liquid Fusarium proliferatum inoculum was the highest,reaching104.7μg·g~(-1), followed by the sample instilled with liquid Hypocrea jecorina inoculum(33.38μg·g~(-1)), and the sample instilled with liquid Hypocrea jecorina inoculum was very low,only1.94μg·g~(-1). It had been shown that the fungus species had great effect on benzyl alcoholcontent, namely, Rhizomucor variabilis had little effect on the formation of benzyl acetone, andFusarium proliferatum the best effect.
5. In this study, the effect of liquid inoculant on benzyl acetone content was comparedwith sterile fungal culture solution. The results had shown that there was little differencebetween liquid inoculant and sterile fungal culture solution. It indicated that there was notdirect relationship between agar formation and the fungal mycelium, on the contrary, there wasclose relationship between agar formation and mycelium metabolites.
6. After instilled with the culture solution of Fusarium proliferatum, on the6thmonthhot-water extractives content in the agar wood was higher than that in no-agar wood by26.923%, which indicated that the cell wall of the timber tissue had been partially damaged andthe metabolites increased; Benzene-ethanol extractives in the agar wood was higher than that inno-agar wood by184.665%, which showed that a part of carbohydrate had been degradated,terpenoid compounds, aromatic compounds content increased accordingly; Holocellulosecontent in the agar wood was lower than that in no-agar wood by21.737%, pentosan(hemicellulose) also lower by9.986%, and lignose content higher by38.316%, which indicatedthat the culture solution could degradate Holocellulose, but not degradate lignose; Comparedwith the no-agar wood, mineral elements content, except for S(increased by24.85%), in theagar wood decreased by7.92%~55.96%, which showed that the timber tissue was under stress, the S content increasing may be the result of passive absorption for self-protection; In theprocess of agar formation, normal balance between growth and respiration had been striked,which resulted in starch conversing to sugar, so the soluble sugar content in the agarwoodincreased by56.10%and starch content decreased by62.92%.
7. Agar material occurred mainly in parenchyma cell of the xylem tracheal and xylem rays.In paraffin sections, no fungi were found where abundant agar engendered, and agar was littlewhere the mycelium was found. Moreover, the fungus found in paraffin sections was notFusarium proliferatum. It had been shown that agar was not metabolites of growth andreproduction of Fusarium proliferatum in timber tissue of Aquilaria sinensis (Lour.) Gilg.
8. After Added Aquilaria sinensis (Lour.) Gilg tissue culture seedling with culture solutionof Fusarium proliferatum and Hypocrea jecorina, there was significant difference in cellmembrane lipid oxidation, cell membrane permeability and glucose metabolism between thetwo groups of samples. From the view of cell membrane lipid peroxidation, the MDA contentin leaves of the samples added Hypocrea jecorina culture solution (H-group) increasedsignificantly, while the one of the samples added Fusarium proliferatum culture solution(F-group) increased less, especially in early phase (24h~48h), which showed that Fusariumproliferatum, as a fungus more conducive to agar formation, could cause Aquilaria sinensis(Lour.) Gilg physiologic stress reaction, but the cell membrane lipid oxidation was not asserious as Hypocrea jecorina. From the view of cell membrane permeability changes, REC(Relative Electrical Conductivity) in the leaves of H-group increased steadily, while F-group sREC increased rapidly at first and then increases slowly. which showed that culture solution ofFusarium proliferatum would damage leaves s cell membrane gradually, while culturesolution of Hypocrea jecorina would damage acutely; From the view of soluble sugar, theleaves’ soluble-sugar in the two groups of samples all changed little in24h, and then allincreased rapidly with H-group more quickly, at the48thhour soluble-sugar content in theH-group samples was significantly higher than that in F-group samples, after the72thhour, thesoluble-sugar content in H-group samples declined more than that in F-group samples, whichindicated that Hypocrea jecorina infection caused nutrient transport disorder and resulted in irregular change of glucose metabolism, while Fusarium proliferatum effect on glucosemetabolism relatively stable.
9. After Added Aquilaria sinensis (Lour.) Gilg tissue culture seedling with culture solutionof Fusarium proliferatum and Hypocrea jecorina, ROS surge phenomenon all occurred in ashort period (0~24h). Later, at the96thhour the superoxide radical and hydrogen peroxide inthe H-group samples were higher than that of F-group samples by4.42U·g~(-1)and26.55U·g~(-1).There were significant differences in active-oxygen scavenging enzymes during thephysiologic stress phase caused by Fusarium proliferatum and Hypocrea jecorina. Thedifferences were manifested in two aspects：firstly, the enzymes activity (including SOD, PODand CAT) of F-group samples were higher than that of H-group samples；Secondly, changeregularity of the enzymes activity was different, enzymes activity in F-group samples changedrelative steadily, while the one in H-group changed acutely.
10. In this study, the hypothesis “physiological stress causing agar” was brought forward.Based on the hypothesis, a trial about agar revulsant formula, including Fusarium proliferatumculture solution, NaCl and JAs, was carried out according to uniform design. The results hadbeen shown that NaCl had greater effect on agarwood quanlity than JAs; the best agarrevulsant formula was Fusarium proliferatum culture solution added1.8248%NaCl and0.0046%JAs. Instilling the trunk with the agar revulsant formula, one year later, maximizedalcohol-soluble extractive content in the wood would be13.6012%theoretically.
引文
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