987 resultados para Bacteriophage T4
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花椒(Zanthoxylum piperitum)是川西干旱河谷地区的重要经济作物,化感作用是花椒连作障碍的原因之一。系统研究花椒化感作用有助于深入理解并最终解决花椒连作障碍。本文通过研究花椒叶、林下土壤浸提液及单一纯化感物质对花椒幼苗生长、苜蓿种子萌发及幼苗生长的影响作用,从生理生化角度揭示浸提液及纯化感物质的作用方式。通过室外和室内模拟实验,对浸提液及纯化感物质的化感效应进行比较,为花椒连作障碍的解决和化感作用机制的深入理解提供依据。主要结果如下: 1.花椒叶及林下土壤浸提液对地下生物量影响作用强与对地上生物量的化感效应,两种浸提液的化感效应强度不同,叶浸提液作用表现更显著。其中在Y6、Y8 、T6和T8处理时,花椒幼苗地下生物量分别降低了31.2%、32.1%、31.6%和31.7%。 2.两种浸提液均能显著影响花椒幼苗体内的保护性酶活性,总体说来,在高浓度下抑制各种抗氧化物酶活性,幼苗体内丙二醛含量增加,幼苗受害严重;在较低浓度下,各种保护性酶活性有所增加,丙二醛含量减少,幼苗伤害减轻。同时,不同月份里,各种酶的活性高低显著不同,9月份的活性显著低于7月份的酶活性。对于养分含量的影响,Y8、T8的影响强度最大,分别使碳元素含量降低了27.8%和30.8%,使钾元素含量降低了34.7%和25.6%。 3.花椒叶及林下土壤浸提液对苜蓿种子萌发及幼苗生长有化感作用,表现在最终萌发率、不同物质代谢及保护性酶活性的差异上。两种浸提液对苜蓿种子萌发过程中蛋白质的含量均无显著性影响,对淀粉和可溶性糖的影响作用类似,高浓度处理无明显化感效应,较低浓度处理显著降低二者在萌发苜蓿种子中的含量。Y2、Y4与T4处理分别使可溶性糖含量减少了32.3%、29.1%和18.8%,Y2与T2处理分别使淀粉含量降低了29.3%和26.8%。 4.苜蓿种子在4种单一化感物质最高浓度即10-3 mol•L-1处理下,萌发率显著降低,半数萌发时间推迟,随着处理浓度降低,抑制作用逐渐减弱,当降低到10-6 mol•L-1时,又能够表现出对苜蓿种子萌发的促进作用。 5.纯化感物质在10-6 mol•L-1时使苜蓿幼苗叶片的保护性酶活性显著升高,丙二醛含量显著降低;在10-3 mol•L-1时使苜蓿叶片中保护性酶活性显著降低,丙二醛含量增加,膜脂过氧化程度加重。 Zanthoxylum piperitum is one of the most important cash crops and has been extensively cultivated in Eastern Tibetan Plateau, especially in the fragile dry valley areas. Allelopathic effects could be a reason for Z. piperitum’s continuous cropping impediment. Systemmatically research of the effect of Z. piperitum allelopathy could help to comprehend the continuous cropping impediment. The allelopathic effects on seedlings growth and seed germination of aqueous extracts of Zanthoxylum piperitum and phenolic allelochemicals were studied, and the action mechanism of the two substances was also discussed from physiology. Indoor and outdoor experiments were set to compare the difference between aqueous extracts and pure allelochemicals. The main results showed that: 1. The aqueous extracts of leaf and soil had significant allelopathic effects on aboveground and underground biomass, but the effect on underground biomass was stronger than the effect on underground evidently. Treated with Y6、Y8 、T6 and T8, the underground biomass was reduced 31.2%、32.1%、31.6% and 31.7% respectively. 2. The activity of activities of superoxide dismutase, catalase, peroxidase and ascorbate peroxidase were significantly reduced, while the content of MDA was increased and the seedlings were suffered stronger, when treated by the high concentration; but at the low concentration, these were reversed. And then, at the different month, the activities of antioxidant enzyme were significantly distinct. As for the contents of nutrient element, Y8、T8 had the more intensive effects than other treatments. 3. The results showed that the two types of aqueous extracts had significant allelopathic effects on seed germination, substances metabolize and the activity of antioxidant enzyme. But the aqueous extracts had no effects on the content of protein, while had the similar effects on the content of starch and soluble sugar. At Y2、Y4 and T4, the content of soluble sugar decreased 32.3%、29.1% and 18.8% respectively. 4. Treated with 10-3 mol•L-1 of the four allelochemicals, the seed germination of alfalfa was significantly inhibited. Ferulic acid, coumarin and vanillic acid at 10-3 mol•L-1 significantly reduced the activities of antioxidant enzyme, while the content of MDA in alfalfa seedling was significantly increased. The restrain effects became weakened with the treat concentration falled. However, ferulic acid, coumarin and vanillic acid could increase the activities of antioxidant enzyme at 10-6mol•L-1.
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利用100MeV/uC离子束对高产酒精酵母菌株进行了辐照诱变的研究。采用红四氮唑作为筛选指示剂,得到了5株产酒能力有所提高的突变酵母菌。利用甜高粱汁的发酵结果表明,T4突变菌株的产酒精能力比原始出发菌株提高了18.6%,且发酵液中的残糖含量也有所降低。随后对T4菌株在甜高粱汁中的最适宜发酵条件做了初步探索,结果表明:最适发酵温度和pH值分别为30℃和4.5。通过10l发酵罐的验证试验表明:在同样发酵条件下,T4菌株的发酵率和产酒精能力都比原始出发菌株提高了12%。
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为了选育出适合发酵甜高粱汁来生产酒精的酵母菌株,本论文以酒精酵母Saccharomyces cerevisiae YY为材料,利用兰州近代物理研究所重离子研究装置(HIRFL)产生的100MeV/u碳离子束对酒精酵母进行了辐照诱变。采用红四氮唑(TTC)作为筛选指示剂,初筛得到了5株产酒能力有所提高的突变酵母菌。通过甜高粱汁发酵,测定发酵液中酒精含量和残糖,复筛出产酒精能力比出发菌株有明显提高的诱变菌株T4。并对其发酵条件进行了优化,以期获得的结果能够为甜高粱汁工业化生产酒精提供参考数据。通过本论文的研究,得到以下初步结果: 1. 在甜高粱汁培养基中,酒精酵母YY的对数生长期在8-20h之间,此时菌体的生长繁殖比较旺盛,活力最佳,为辐照诱变的最佳时期。辐照后,菌体的存活率随辐照剂量的增加呈现出逐渐衰减的趋势。 2. 红四氮唑TTC是一种无色显色指示剂,活菌中所含的脱氢酶可将它还原成红色,因此可以根据菌落呈色的深浅判断酵母菌产酒精能力的高低,从而挑选出产酒能力较高的菌株。本试验用TTC双层培养基法初步筛选出了利用甜高粱汁发酵生产酒精能力较强的T4酵母菌株。 3. 对影响T4菌发酵甜高粱汁生产酒精的几个主要因素(甜高粱汁糖度、接种量、温度、pH、无机盐)进行了初步探讨研究,得出了T4菌发酵甜高粱汁生产酒精的最适条件为:甜高粱汁糖度22%,接种量10%,温度30oC,pH 4.5 ,无机盐加入量为:(NH4)2SO4 1g/L,KH2PO4 5g/L,MgSO4 3g/L。 4. 对发酵条件进行优化后的中试结果显示:出发菌株YY发酵甜高粱汁的时间为36h,酒精产量为8.6% (V/V) ,而T4突变菌甜高粱汁发酵液中的最终酒精含量可以达到9.8%,发酵时间仅为24h。因此,T4菌在工业应用中很有前景
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In bacteriophage, transcription elongation is regulated by the N protein, which binds a nascent mRNA hairpin ( termed boxB) and enables RNA polymerase to read through distal terminators. We have examined the structure, energetics and in vivo function of a number of N boxB complexes derived from in vitro protein selection. Trp18 fully stacks on the RNA loop in the wild-type structure, and can become partially or completely unstacked when the sequence context is changed three or four residues away, resulting in a recognition interface in which the best binding residues depend on the sequence context. Notably, in vivo antitermination activity correlates with the presence of a stacked aromatic residue at position 18, but not with N boxB binding affinity. Our work demonstrates that RNA polymerase responds to subtle conformational changes in cis-acting regulatory complexes and that approximation of components is not sufficient to generate a fully functional transcription switch.
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Iodothyronine plays a major role in growth, basic metabolism and organ formation. It has an extremely limited source in the body. In this thesis, we designed iodothyronine(T4) as hapten. Then a single chain antibody displayed on phange was obtained from a human phage displaying a single chain antibody library. The specific genes of E3 was subcloned in P-5E vector. According to its amino acid sequences, we simulate its three dimention structure by computer. It has never been reported in PDB.
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化合物的结构与其活性具有相关性 .建立相关性需要由结构提取特征 ,并运用这些特征作为自变量去构造数学模型 ,以便预测未知的化合物及其活性 .目前 ,在诸多 3D- QSAR方法中 ,应用最为广泛的是 Co MFA( Comparasive Molecular Field Analysis)方法 [1~ 3] ,但新近的研究发现 ,Co MFA方法对于某类化合物 (如苯胺类化合物 )适应性较差 ,但分子在三维空间投影面积及量化参数的引入可大大改善所得结果[4 ] .本文从另一角度 (即广义二面角 )研究了分子在三维空间的形状 ,得到了满意的结果 .1 实验部分本文选用 34个 1 - [( 2 - Hydroxyethoxy) methyl]- 6- ( phenylthio) thymine( HEPT)类化合物 [5] ,通式见Scheme 1 HEPT derivative substituted HEPTScheme 1 .表 1中分别列出了 34个分子对应的 R,X,Y基团 .该类化合物是具有较高生物活性的抗艾滋病药物 ,其活性以 EC50 表示 ( EC50 为降低 5 0 %T4细胞而被 HIV...
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甲状腺素脱碘酶是能够催化甲状腺激素不同降解反应的一簇含硒酶 [1,2 ] ,由于其含量低 ,分离纯化困难等[3] ,严重阻碍了其生物化学性质和应用研究 .自 Lerner[4 ] 和 Schultz等 [5] 分别制备出了具有催化作用的抗体酶以来 ,抗体酶在天然酶的模拟研究中已展示了巨大潜力 ,有些抗体酶已达到或超过了天然酶的活力 [6 ] .我们曾用抗体酶技术制得具有脱碘酶活性的抗体酶 [7] .为进一步提高脱碘抗体酶活力 ,本文以疏水腔修饰法 [8] 为指导 ,用铜离子络合保护法设计合成了 3种甲状腺素衍生物半抗原和全抗原 ,并制备出了一种半抗原的单克隆抗体 .试剂和仪器 :五水甲状腺素钠 ( T4 )为 SIGMA产品 ;牛血清白蛋白 ( BSA )和卵清白蛋白( Ovalbumin) ,SIGMA分装 ;戊二醛 ,硫酸二甲酯 ,溴化苄 ,对硝基溴苄和其它试剂均为国产分析纯 .Finni- gan mat LCQ电喷雾多极串联质谱仪 ;KONTRON公司 92 2型紫外可见分光光度仪 ;德国Elementar Anallysensysteme Gmb H Vario EL元素分析仪 ;美国 Varian...
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Conversion of thyroxine (T-4) to 3,5,3'-triiodothyronine is an essential first step in controlling thyroid hormone action. Type I deiodinase (DI) can catalyze the conversion to produce the bulk of serum 3,5,3'-triiodothyronine. Acting as a mimic of DI, a selenium-containing catalytic antibody (Se-4C5) prepared by converting the serine residues of monoclonal antibody 4C5 raised against T4 into selenocysteines, can catalyze the deiodination of T4 with dithiothreitol (DTT) as cosubstrate. The mimic enzyme Se-4C5 exhibited a much greater deiodinase activity than model compound ebselen and another selenium-containing antibody Se-Hp4 against GSH. The coupling of selenocysteine with the combining pocket of antibody 4C5 endowed Se-4C5 with enzymatic activity. To probe the catalytic mechanism of the catalytic antibody, detailed kinetic studies were carried out in this paper. Investigations into the deiodinative reaction revealed the relationship between the initial velocity and substrate concentration. The characteristic parallel Dalziel plots demonstrated that Se-4C5-catalyzed reaction mechanism was ping-pong one, involving at least one covalent enzyme intermediate. The kinetic properties of the catalytic antibody were similar to those of DI, with K-m values for T-4 and DTT of approximately 0.8 muM and 1.8 muM, respectively, and a V-m value of 270 pmol per mg of protein per min. The activity could be sensitively inhibited by 6-propyl-2-thiouracil (PTU) with a K-i value of similar to 120 muM at 2.0 muM T-4 concentration. The PTU inhibition was progressively alleviated with the increasing concentration of added DTT, revealing that PTU was a competitive inhibitor for DTT.
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Vibrio harveyi is an important marine pathogen that can infect a number of aquaculture species. V. harveyi degQ (degQ(Vh)), the gene encoding a DegQ homologue, was cloned from T4, a pathogenic V. harveyi strain isolated from diseased fish. DegQ(Vh) was closely related to the HtrA family members identified in other Vibrio species and could complement the temperature-sensitive phenotype of an Escherichia coli strain defective in degP. Expression of degQVh in T4 was modulated by temperature, possibly through the sigma(E)-like factor. Enzymatic analyses demonstrated that the recombinant DegQVh protein expressed in and purified from E. coli was an active serine protease whose activity required the integrity of the catalytic site and the PDZ domains. The optimal temperature and pH of the recombinant DegQVh protein were 50 C and pH 8.0. A vaccination study indicated that the purified recombinant DegQVh was a protective immunogen that could confer protection upon fish against infection by V. harveyi. In order to improve the efficiency of DegQVh as a vaccine, a genetic construct in the form of the plasmid pAQ1 was built, in which the DNA encoding the processed DegQVh protein was fused with the DNA encoding the secretion region of AgaV, an extracellular beta-agarase. The E.coli strain harboring pAQ1 could express and secrete the chimeric DegQVh protein into the culture supernatant. Vaccination of fish with viable E. coli expressing chimeric degQ(Vh) significantly (P < 0.001) enhanced the survival of fish against V. harveyi challenge, which was possibly due to the relatively prolonged exposure of the immune system to the recombinant antigen produced constitutively, albeit at a gradually decreasing level, by the carrier strain.
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Vibrio harveyi is an important marine pathogen that can infect a number of aquaculture species. V. harveyi degQ (degQ(Vh)), the gene encoding a DegQ homologue, was cloned from T4, a pathogenic V. harveyi strain isolated from diseased fish. DegQ(Vh) was closely related to the HtrA family members identified in other Vibrio species and could complement the temperature-sensitive phenotype of an Escherichia coli strain defective in degP. Expression of degQVh in T4 was modulated by temperature, possibly through the sigma(E)-like factor. Enzymatic analyses demonstrated that the recombinant DegQVh protein expressed in and purified from E. coli was an active serine protease whose activity required the integrity of the catalytic site and the PDZ domains. The optimal temperature and pH of the recombinant DegQVh protein were 50 C and pH 8.0. A vaccination study indicated that the purified recombinant DegQVh was a protective immunogen that could confer protection upon fish against infection by V. harveyi. In order to improve the efficiency of DegQVh as a vaccine, a genetic construct in the form of the plasmid pAQ1 was built, in which the DNA encoding the processed DegQVh protein was fused with the DNA encoding the secretion region of AgaV, an extracellular beta-agarase. The E.coli strain harboring pAQ1 could express and secrete the chimeric DegQVh protein into the culture supernatant. Vaccination of fish with viable E. coli expressing chimeric degQ(Vh) significantly (P < 0.001) enhanced the survival of fish against V. harveyi challenge, which was possibly due to the relatively prolonged exposure of the immune system to the recombinant antigen produced constitutively, albeit at a gradually decreasing level, by the carrier strain.
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VhhP2 is an Outer membrane protein identified in a pathogenic Vibrio harveyi strain, T4, isolated from diseased fish. When used as a Subunit Vaccine, purified recombinant VhhP2 affords high level of protection upon Japanese flounder against V harveyi challenge. Vaccination with VhhP2 induced the expression of a number of immune-related genes, especially those encoding immunoglobulin M (IgM) and major histocompatibility complex (MHC) II alpha. A VhhP2 surface display system, in the form of the fish commensal strain FIR harboring the vhhP2-expressing plasmid pJVP, was constructed. PF3/pJVP is able to produce and present recombinant VhhP2 on cell surface. Vaccination of fish with live PF3/pJVP via intraperitoneal injection elicited Strong immunoprotection. Vaccination of fish orally with live PF3/pJVP embedded in alginate microspheres also induced effective immunoprotection. In addition, a VhhP2-based surface display system was created, in which VhhP2 serves as a carrier for the Surface delivery of a heterologous Edwardsiella tarda immunogen, Et18, that is fused in-frame to VhhP2. DH5 alpha/pJVP18, which expresses and surface-displays the VhhP2-Et18 chimera, proved to be an effective vaccine that call protect fish against infections by V. harveyi and E. tarda to the extents comparable to those produced by vaccination with purified recombinant VhhP2 and Et18, respectively. These data suggest that VhhP2 may be applied as a vaccine and a vaccine carrier against infections by V. harveyi and other pathogens such as F. tarda. (C) 2009 Elsevier Ltd. All rights reserved.
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以奥利亚罗非鱼(Oreochromis aureus)为实验对象,设计了3种不同的摄食类型,分别是鲜活饵料组、饥饿3周后饱食投喂组和人工饲料组。鲜活饵料组投喂冰冻赤子爱胜蚓,利用蚯蚓体内丰富的营养成分和活性物质,以期获得奥利亚罗非鱼良好的生长状况;饥饿后饱食组是指饥饿3周后,以人工饲料饱食投喂2周,用于研究饥饿与补偿生长获得快速生长时血液理化指标的变化情况;人工饲料组作为对照组。纯淡水条件下养殖,水温25±2℃。测定了奥利亚罗非鱼在3种摄食类型饲喂下某些血液生理生化指标变化的情况,并将指标变化情况与增重率做相关性分析,试图找出能够反映奥利亚罗非鱼生长性能的血液生理生化指标。 研究结果表明,奥利亚罗非鱼在饥饿3周后获得了补偿生长,补偿生长时的增重率和特定生长率显著高于人工饲料组(P<0.05),高于鲜活饵料组,但差别不显著;相关性分析研究表明血清总蛋白、胆固醇、四碘甲状腺原氨酸(T4)与增重率极显著相关(P<0.01),血红蛋白显著相关(P<0.05),红细胞、白细胞、碱性磷酸酶高度相关(相关系数为0.580、0.551和0.557),因此,建议血清总蛋白、胆固醇和血红蛋白可作为能够反映罗非鱼生长性能的新指标。 根据序列设计引物,PCR反应条件:变性温度:95 ℃,3 min;退火温度:57℃,20 sec;延伸温度:72℃,5 min,共36个循环,从牙鲆、黑鲪和鲈鱼中克隆出胰岛素样生长因子(IGF-Ⅰ)部分序列,首次证实了IGF-Ⅰ在3种海水鱼中的存在。 利用蛋氨酸与ZnSO4•7H2O,在pH 5.5、80℃下,反应1小时,采用蛋氨酸与硫酸锌2:1的配料比,合成出了产物蛋氨酸螯合锌,蛋氨酸螯合锌外观白色,粉状,室温下微溶于水,不溶于乙醇,并用原子吸收光谱法测定其含锌量为15%,螯合率为88.2%。
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从山东黄岛海水养殖场分离到一株弧菌V134,从中克隆得到琼胶酶基因agaV,并将其在大肠杆菌中表达,纯化得到重组的琼胶酶AgaV。酶活分析发现该琼胶酶的最适温度在40℃左右,对pH比较敏感,pH 7.0时具有最高的琼胶裂解活性。对AgaV进行了两种应用性探索:(1)利用AgaV从琼脂糖凝胶中回收DNA,回收效率可达90%以上;(2)利用agaV作为报告基因构建了捕获分泌序列的载体pBU,并用其从革兰氏阳性细菌(G+菌)和革兰氏阴性细菌(G-菌)中筛选出了一系列分泌蛋白。将利用pBU从一株哈维氏弧菌T4中筛选出的6个分泌蛋白分别进行基因克隆、蛋白表达纯化和牙鲆免疫实验,发现其中一个蛋白,命名为DegQVh,具有免疫保护效应,其免疫保护率(RPS)可达64%。为了提高DegQVh的免疫保护效应,将AgaV的分泌结构域与DegQVh融合,构成融合抗原AgaV-DegQVh。利用大肠杆菌作为载体菌构建了AgaV-DegQVh融合抗原递呈系统,用其作为疫苗进行免疫,发现其RPS可达到95%。酶活分析表明DegQVh在50℃、pH 8.0时具有最高的活性。突变分析表明83位的组氨酸、113位的天冬氨酸和188位的丝氨酸以及两个PDZ结构域是DegQVh活性所必需的。表达分析发现degQVh表达受温度和细胞浓度调控,并且其上游有一个受E调控的启动子。进一步的分析发现DegQVh能够与大肠杆菌的DegP功能互补。
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测定分析了新疆地区引入澳大利亚萨福克羊及无角陶塞特羊的体温、脉搏、呼吸频率、血清T3、T4 含量、繁殖率、总适应能力及杂一代公羔的育肥屠宰性能 .结果表明 ,萨福克羊及无角陶塞特羊繁殖率及体重接近或超过原产地澳大利亚的生产水平 .萨福克羊总适应能力为 98.80 % ,无角陶赛特羊为 98.74% .杂一代公羔日增重、饲料转化率、净肉重较哈萨克羊高 ,而尾脂重低于哈萨克羊。
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本项研究以长江中下游地区的浅水湖泊(太湖、巢湖、龙感湖)为研究对象,碳、氮同位素为研究手段,结合210Pb和14C年代学,以及沉积物中TOC、TN、C/N比值、TP等多种地球化学参数,对近代沉积环境演化过程,沉积物有机质来源以及西太湖形成演化的古环境进行分析研究。通过研究,得到以下主要认识: 1. 太湖沉积物剖面上δ13Corg和C/N比值分布范围指示太湖沉积物的有机物质来源主要是水生藻类。竺山湾T3点基本没有陆源物质输入;梅梁湾T2点有部分陆源物质输入;湖心T4点沉积物有机质来源还可能是不同于梅梁湾和竺山湾的水生植物。沉积物的地球化学参数剖面指示太湖从1920s始,沉积环境受人为因素的影响而逐渐营养化。 2. 巢湖沉积物δ13Corg和δ15N的研究结果,不仅指示沉积物有机质来源主要是水生藻类,受城市污染和农业面源污染输入的一定影响,而且指示了在沉积历史上因为巢湖闸的建成,湖泊生产力和浮游植物物种也发生了改变,湖泊富营养化。 3. 龙感湖沉积物δ13Corg和δ15N的研究分析,表明湖泊有机沉积以自身有机物源为主,受陆源输入影响小。1960s围垦造成营养盐输入量的增加,因为草型湖泊这一特征有利于营养盐的积累,缓解水体的富营养化程度,龙感湖始终处于中营养程度。 4. 太湖、巢湖和龙感湖表层沉积物孔隙水中NO3-和SO42-含量的差异,主要受水域污染状况、底泥有机质的丰富情况、环境水动力条件的变化、泥沙沉积和再悬浮过程引起的元素累积和释放影响,也是藻型湖泊与草型湖泊生产力大小、有机质沉积通量以及微生物丰度和活性等差异的表征。 5. 沉积物表层吸附态NH4+-N含量,体现了水域环境受污染的严重程度是:龙感湖<巢湖<太湖梅梁湾。三个湖泊中底泥有机质丰度差异和生物参与的氨化作用差别都非常显著。三个湖泊的沉积物表层铵态氮的含量,均表明了沉积物表层向上覆水体可能具有潜在的铵态氮迁移趋势。 6. 太湖古环境分析研究中,竺山湾和梅梁湾沉积物剖面的粒度分析、δ13Corg、δ15N、TOC、TN、C/N比值和TP都随沉积深度,对应14C定年结果,指示了西太湖沉积演化历程的三个阶段及其有机质来源。6870~6670 a B.P.,研究区被咸水覆盖,有机质来源是典型的水体自生来源,在竺山湾有逐渐增加的C4植物的输入。6670~5140 a B.P.,可能形成瀉湖并出现沉积间断。5140 a B.P.至今,形成淡水湖泊,沉积有机质主要来自湖泊自生物源,存在沉积间断。 7. 长江中下游的浅水湖泊沉积物中的有机质来源主要以湖泊自生来源为主,因为水体初级生产力的升高受陆源物质输入增加的影响,流域内人类活动引起的陆源物质输入不容忽视。长江中下游湖泊的近代沉积环境演化过程因为近岸距离、水动力强度和发育不同的水生植物等因素存在区域性差异,导致湖泊富营养化的最大根源是人为因素的影响发生的环境演变。