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_<_<_<_<_<<<_<_<_<_<_<_<_<_<_<___<___<___<___<_______<__________________________________<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<_____<_<_<_<_<_<___<___<______________________________________________________<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<<_<<<_<<<_<<<_<<<_<<<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<___<___<___<_______<_______<____________<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<____________________________________________________________________________________<<<<<<<<<<<<<_<<<<<<<_<<<_<<<_<<<_<<<_<<<_<<<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<___<_<_<___<_<_<___<___<_____________________________<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<_<_<___<______________________________________________________________________________<<<<<<<<<<<<<<<_<<<_<<<_<<<_<<<_<<<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<___<___<_<_<_<_<___<___<___<___<___<_______<________<_<_<_<_<_<<<<<<<<<_<_<_<_<_<_<_<___<___<___<___<________________________________________________________________________________<_<<<<<<<_<<<_<<<_<<<_<<<_<_<_<_<<<<<<<<<<<<<<<<<<<<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<___<___<___________________<___<__________________________<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<<<<<<<<<<<<<<<<<<<_<_<_<_<___<___<______________________________________________________________________________<<<<<<<<<<<<<<<<<<<_<<<_<<<_<<<_<_<_<_<_<_<_<_<_<_<_<_<<<<<<<<<<<<<<<<<_<_<_<_<_<_<_<_<_<_<_<_<_<___<_<_<___<___<___<___<___<_______<_______<_______<____________<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<_____________________<<<_<_<_<_<____________________________________________________________________________________<<<<<<<<<<<<<_<<<_<<<_<<<_<<<_<_<_<_<_<_<_<_<_<_<_<___<___<___<___<_<<<<<<<<<_<<<_<_<_<_<_<_<_<_<_<___<___<___<___<_______<______________________________________<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<__________________________<<_<_<_<_<______________________________________________________________________________<<<<<<<<<<<<<<<<<<<<<<<_<<<_<<<_<_<_<_<_<_<_<_<_<_<_<_<_<___<___<___<___<_______<<<_<<<_<_<_<_<_<_<_<___<___<___<___<___<___<___<_______<___________<___<_______--&TNPP &՜.+,0     屏幕显示gNB ;h oTimes New Roman宋体 华文新魏楷体_GB2312PulseMicrosoft 公式 3.0 画笔图片G第三节 代谢网络中碳架 物质流的调动 处于一定环境条件下的微生物培养物中,参与代谢的物质在代谢途径(网络)中按一定规律流动,形成微生物代谢的物质流。j 代谢物质的流动过程是一种类似“流体流动” 的过程,它具备流动的一切属性,诸如 方向性、连续性、有序性、可调性等等, 并且可以接受疏导、阻塞、分流、汇流等 “ 治理 ”, 也可能发生“ 干枯 ”和“溢出 ( 泛滥 ) ”等现象。此外,还有流速和流量等问题。 化能异养型微生物以有机化合物为碳源和能源。因此,在讨论化能异养型微生物代谢流动时,首先抓住碳元素代谢这个主要矛盾,研究微生物细胞的碳架物质流。{4.3.1 碳架物质在代谢途径(代 谢网络)中的流动 4.3.2 微生物的代谢流及其治理 !4.3.1 碳架物质在代谢途径(代谢网络)中的流动 在典型的工业发酵过程中,微生物的代谢流包括营养物质进入细胞、代谢中间物(碳架物质)在代谢途径中的流动和代谢产物从细胞进入环境这样顺序发生的过程。n4.3.1.1 有机化合物的组成元素及它们 各自在生命活动中的作用(略) 4.3.1.2 碳架物质在代谢途径中的流动 以代谢流动的观点加深对代谢网络的认识:碳架物质在网络的有关途径中流动,从原料到终端产物碳架物质流经的路径叫做载流路径。 ?4.3.1.3 典型工业发酵的工作模式 典型的工业发酵要从发酵液中获得微生物的代谢产物,也就是要让原料有机化合物在微生物细胞的有关的代谢途径(载流路径)中流动的过程中,转化成工业发酵的目的产物,最后目的产物分泌出细胞。  上图中,白色椭圆代表细胞的界面,界面包围部分是细胞机器,界面外侧是细胞机器所在的环境。带箭头的黑线代表典型的工业发酵的细胞机器的载流路径。 N代表营养物质, P代表目的产物。)4.3.2 微生物的代谢流及其治理P 我们曾从热力学的角度把微生物看作远离平衡状态的不平衡的开放体系;我们曾借助电学的知识来讨论生物氧化和代谢能的问题;我们曾借用互联网的概念来讨论代谢网络的延伸问题;在这里我们要借用流体力学的常识来分析微生物细胞的代谢流。  如果把碳架物质流看作水一样的流体,那么按照工业发酵的工作模式,就可把微生物活细胞看作是有一个总的进水口和一个总的出水口的、包含特定的管道网的大容器,组成管道网的管道有粗有细,管道网的管道有分支、有交叉,有可控的阀门。代谢物质流就如同自来水一样在管道和阀门的控制下流进和流出细胞,形成代谢流。U 在这个认识的基础上,结合代谢能支撑假说和代谢网络假说的内容,讨论微生物的代谢及代谢流的变动性;结合本章( 代谢调节的 3 个子系统 )的内容,讨论代谢流对环境条件的响应和代谢流治理的可能性。最后还要讨论有关代谢节点的刚性的问题。] 微生物的代谢途经的变动性和可调节性,不但是代谢流飘移的内在原因,而且也是人工控制代谢流的生物学根据。工业发酵中,人的主观能动作用表现在对代谢流的治理上。治理的直接的目的是让微生物细胞过量合成( overproduction )人们所需要的代谢产物。4.3.2.1 碳架物质在代谢网络中流经的 分支处及各分支的流量分配 4.3.2.2 过量合成与微生物的异常代谢 4.3.2.3 溢出代谢 4.3.2.4 代谢流治理的可能性和现实性 4.3.2.5 代谢网络的刚性4.3.2.1 碳架物质在代谢网络中流经的分支处及各分支的流量分配 我们把代谢网络中代谢途径的交叉点叫做代谢网络的 “ 节点 ”, 实际上它们是碳架物质在代谢网络中流经的代谢中间物。初级代谢网络的中心板块上的 12 种前体代谢物是当然的节点,而且还是代谢途径的分流或汇流处。 o 节点处不论发生汇流还是分流,都存在流量分配的问题。这个分配既与遗传因素有关,又与条件因素有关。不同微生物物种的代谢途径基本一致,因此,可在微生物学有关工具书查到有关的途径;而代谢调节机制并不完全一样,不一致的情况可在有关论文或综述中找到。 D 在生物化学中,早有一套描述代谢途径的调节机制的方法,例如,用一个从效应物(途径的终端产物)引出的、指向途径中某个代表酶的短箭头的长箭头来标示反馈调节。这种标示方法对于线状途径或偶有分支的途径来说,是很有用的,但是,如果在代谢网络图中使用,代表调节的箭头与代表酶的箭头交织在一起,容易引起误解。 因此我们要启用一种新的代谢调节图图例,用来标记代谢网络图。这种标记方法有四个要点: 6① 标明调节方式 用“I”表示抑制(inhibition),用“A”表示激活( activation ),用 “R” 表示阻遏(repression); ②标明效应物 一般用三个字母标示效应物( 调节特定的酶的水平或活性的化合物 叫做这种酶的效应物) ;PowerPoint 演示文稿 例如:某效应物名称后面加上 “I”, 标在代表酶的箭头旁边,表示该效应物对该酶的抑制作用;效应物名称后面加上 “A”,标在代表酶的箭头旁边,表示该效应物对该酶的激活作用;效应物名称后面加上 “R”,标在代表酶的箭头旁边,表示该效应物对该酶的阻遏作用。如果 2 种效应物协同作用,则在两个效应物的名称都写上,再在后面加上对应的英文字母。 PowerPoint 演示文稿 上图(碳架物质在 EMP和 HMP途径上的分流控制图)是一张按此方法标示的网络图,请按此方法读图,并从以下说明中得到帮助。 大肠杆菌细胞磷酸果糖激酶(PFK)的酶水平随条件而变动,有氧生长中细胞的PFK水平低于厌氧生长中的细胞,有氧生长中的细胞的PFK(由二聚体组成),对ATP不敏感,厌氧生长中的细胞的PFK (由四聚体组成),对ATP敏感。  在有氧条件下,由于PFK的低水平和 FDP 醛缩酶(FDPA)的极端低水平,大肠杆菌主要经 HMP 途径降解葡萄糖。因为在有氧条件下 PFK 对 ATP不敏感,且其活力高于FDP醛缩酶(FDPA ),FDP 有累积,所以它可能对 HMP 途径的 6-磷酸葡萄糖酸脱氢酶( 6PGDH )起抑制物的作用,使6PGDH成为HMP降解流上的一个“阀门”(有氧条件下的控制“阀门”)。 在厌氧条件下,PFK和FDPA水平较有氧时高,因而 EMP 途径对葡萄糖降解代谢的参与程度增加。 因为 PFK 的水平仍高于FDPA,FDP 仍然累积并继续抑制 6PGDH,但随着EMP参与程度的增加,EMP 途径所产生的NADH浓度增高。NADH是6-磷酸葡萄糖脱氢酶( G6PDH )的效应物,NADH 浓度上升就会抑制 G6PDH,使 G6PDH 成为 HMP降解流上的另一个 “ 阀门 ” ( 厌氧条件下的控制 “ 阀门 ” )。  这种抑制作用是必要的,因为提高葡萄糖浓度可以使 FDPA水平上升许多倍,而 PFK 水平基本保持不变,这样 FDP 浓度势必是低的,以致无法控制 6PGDH ,这时 NADH就通过对 G6PDH 的抑制作用控制 HMP 降解流。这种两个“阀门 ”交替控制的机制可以解释为什么在严格厌氧条件下,大肠杆菌经 HMP 途径降解的葡萄糖只占葡萄糖降解总量的20%~30%。J4.3.2.2 过量合成和微生物的异常代谢 野生菌株在它们天然栖息地在其生长或维持,需要时也能合成人们所希望的代谢产物,但在其固有的代谢调节机制的控制下合成的量不多不少,只够细胞自身生长或维持的需要。体现了微生物细胞的经济性。 适用于发酵生产的微生物必须具有有过量合成的能力,也就是说是其目的产物的合成量必须大大高于它通常的合成量。要做到这一点,不仅要求合成目的产物的途径畅通,还必须有将目的产物排出细胞的满意的机制。过量合成的速率取决于微生物的遗传性能,以及在培养条件下细胞的生理状态。 把自然界中自然竞争中幸存的微生物称为野生型微生物,它们具有比较严密的代谢调节功能,至少在它们幸存的环境条件下不会过量合成初级代谢产物。在这样的细胞经济体系中,细胞的生产力体现在其自身的增殖。细胞固有的生产关系支持细胞自身的增殖(生产细胞),不支持(人的) 目的产物的过量生产(生产特定的初级代谢产物)。 要解放细胞对于目的产物(特定的初级代谢产物)的生产力,就有必要改变细胞原有的生产关系。对微生物进行遗传改造,就是要修正原有的生产关系。? 微生物过量合成目的产物(特定的代谢中间产物)的过程,实质上是功能残缺或遗传失常的微生物细胞在人为的特定条件下进行异常代谢的过程。因此,研究微生物过量合成指定的初级代谢产物的机理,就是研究工业微生物的病理学。 微生物排出某些能量代谢副产物的过程,实质上是兼性厌氧微生物在环境条件逼迫下(如缺氧),变通地维持生计的过程。兼性厌氧微生物细胞在缺氧的条件下,以变通的方式实现还原力的平衡和辅酶的再生和回用,使细胞的能量代谢得以维持,与此同时形成并分泌出能量代谢副产物。例如酿酒酵母在发酵液中累积能量代谢副产物乙醇的过程,就是这样的过程。4.3.2.3 溢出代谢 发生溢出代谢(overflow metabolism)的原因是,代谢途径前段的运转速率大于后段的运转速率。采用改变环境条件或限制营养供应等方法,可促使微生物进行溢出代谢。 这里介绍两个溢出代谢的实例: ⑴TCA环中间产物柠檬酸的溢出代谢, ⑵导致能量代谢的副产物乙醇生成的溢出代谢。 ⑴ 柠檬酸的溢出代谢: 多种微生物均能因受激而过量合成柠檬酸。研究柠檬酸溢出代谢的最好的例子无疑是黑曲霉。黑曲霉之所以能在特定环境条件下累积柠檬酸,是因为在这种环境条件下代谢途径前段的运转速率大于后段的运转速率。柠檬酸的溢出代谢是黑曲霉特有的遗传和生化机制与培养条件共同起作用的结果。引起溢出代谢的原因包括以下三个方面:PowerPoint 演示文稿N ①高水平的柠檬酸合成能力。这个能力由 3 个因素构成。第一是在有高浓度草酰乙酸(OAA )的情况下对 AcCoA 具有高度亲和力的组成型的柠檬酸合成酶 ( CS ) 的存在;第二是催化丙酮酸 ( PYR )固定 CO2 生成草酰乙酸反应的高水平的组成型的丙酮酸羧化酶( PC ) 的存在;第三是在缺少锰的条件下,蛋白质分解或蛋白质合成受阻造成的铵的高浓度能解除柠檬酸(CTA)对磷酸果糖激酶(PFK)的抑制。此外,柠檬酸的分泌降低其胞内浓度。 ②较低的降解柠檬酸的能力。这能力由两个因素构成。第一是低水平的α-酮戊二酸脱氢酶( KD )影响 TCA环运行的畅通程度, 使TCA环前半部的中间产物积压; 第二,在锰缺乏的条件下,顺乌头酸酶( AE )和 异柠檬酸脱氢酶 (ID)的活性降低,从而使柠檬酸的累积比其它几种酸( 顺乌头酸、异柠檬酸和α-酮戊二酸)更明显。 ③ 在柠檬酸过量合成阶段,培养基的 pH 值显然会影响细胞膜对目的产物柠檬酸和的跨膜输送;柠檬酸的分泌也会影响培养基的 pH 值。锰与铁的缺乏有利于柠檬酸的排出。0⑵ 乙醇的溢出代谢: 葡萄糖为碳源,对酿酒酵母进行批式培养,出现二段生长( two growth phase)现象:在第一阶段,葡萄糖浓度较大,萄萄糖的消耗伴随着乙醇的产生;第二阶段,葡萄糖浓度下降到某值以下,乙醇被氧化。w 这样的二段生长现象曾被认为是呼吸途径的阻遏和解阻遏,而且认为是分解代谢物阻遏(此名称已被营养阻遏代替,见本章4.1.2.1 )。后来才认识到,酿酒酵母之所以在葡萄糖培养基中累积乙醇,是因为该微生物有限的呼吸能力,并把这种现象称为瓶颈效应( bottleneck effect)。j 细胞并不会因为葡萄糖的高速消耗而附带地形成足够高的呼吸能力(指呼吸链使NADH再生的能力)与之配套;而且葡萄糖的异生成途径(需NADH)因受到高浓度的葡萄糖的调节而受阻,更加强了瓶颈效应。因此瓶颈是对生物氧化中呼吸链电子传递通道的相对狭窄而言的。  在供氧情况下,酿酒酵母细胞中葡萄糖主要经 EMP-TCA 途径( 少量经HMP途径 )氧化降解。这些途径运行中形成的NADH和FADH2分子上的电子需经电子传递链(呼吸链)传递给分子氧。 当葡萄糖浓度较低时,葡萄糖氧化降解过程所需配套的电子传递能力处于电子传递链传递能力范围以内,这时氧的吸收速率与细胞的生长速率成线性关系。辅酶NADH 和辅基 FADH2 完全可以在电子传递链释放电子而得到再生。C 当葡萄糖的浓度超过某值,葡萄糖氧化降解过程所需配套的电子传递能力超越微生物细胞电子传递链传递能力范围,一部分NADH不能通过电子传递链而再生成 NAD+, 只好将电子交给内源的有机化合物乙醛,将乙醛还原成乙醇,从而实现还原力的平衡。从而导致生成乙醇的溢出代谢。由此可见,乙醇的溢出代谢实际上是细胞内生物氧化过程中形成的还原力相对于电子传递链的呼吸能力的溢出,推动了乙醛还原生成乙醇的发酵过程。 从以上实例可以看到,溢出代谢的发生,不但要求微生物本身具有相应的遗传根据,还必须具备相应的环境条件。 对于工业发酵来说,就是特定的生产菌种和与其对应的生产工艺条件。4.3.2.4 代谢流“治理”的可能性和 现实性 代谢流具有流体流动的基本属性,因此具有“治理”的可能性;微生物代谢的变动性和微生物遗传的保守性,加上人的主观能动性,能使可能变为现实。 本课程主要讨论初级代谢产物的过量合成,兼顾次级代谢产物的合成。1 现代发酵工程依靠微生物活细胞来完成发酵工厂生产线上不可缺少的加工(或转化)步骤,在这个意义上,微生物活细胞就是工业发酵生产线上不可缺少的微生物生物机器(或称细胞机器),是发酵工厂里关键的生产机器。* 细胞机器与普通意义上的机器不同,它们必须由代谢能 ( metabolic energy)直接驱动。在发酵工厂的生产线上,微生物必须完成微生物细胞机器的在线制造(细胞增殖)和生产运行(目的产物的合成和输出)的双重任务。 根据工业发酵的现状,把依靠化能异养型微生物合成某最终被分泌到胞外的初级代谢产物的过程,作为工业发酵的典型过程来研究,并把代谢研究的重点放在碳的代谢上。从而建立了生物机器(又称细胞机器)工作的基本模式。在此工作模式的基础上,提出了工业发酵中微生物代谢的五段式和初级代谢产物的过量合成的五字策略。 在物竞天择的基础上建立的细胞经济体系已经经受长期的实践检验,并已成为微生物细胞生命活动的保障体系。微生物通过生存竞争而获得的一整套自我调节机制,原本服务于微生物的自然竞争和生命保障,因此它不会大量的合成维持自我生存之外的代谢产物。而人类因自我发展的需要,却希望微生物能够大量地合成能满足人类需求为目的产物(微生物的代谢产物)。 k 微生物代谢产物的生物合成是系统工程,要完成这一类系统工程,必须依靠微生物的高度有序的生命活动,决不是打乱它,而是在可能的范围内对它进行有效的引导。 工业发酵呼唤人与微生物的合作,而在合作中,人处于主导的地位,人应该发挥主观能动作用。%⑴ 初级代谢产物的合成 初级代谢产物是指这样一些低分子质量的合成途径的终产物,它们是构建生物大分子的分子模块,是微生物营养性生长所必需的生长因子,如氨基酸、嘌呤和嘧啶的核苷与核苷酸以及维生素。 初级代谢产物都是从中心代谢途径的中间产物(12种前体代谢物)开始合成的。因为它们都是微生物营养性生长所必需的,除了遗传上有缺陷的菌株外,微生物细胞具有齐全的初级代谢途径基因,也就是说在合适的环境条件下它们的初级代谢途径是普遍存在的。初级途径上的酶的特异性比次级代谢的酶的特异性要高。 = 各种微生物的初级代谢产物的合成途径基本相同,在同一属微生物中,初级代谢产物合成途径的差异是十分罕见的,但在合成的调节机制上有较大的差异。 初级代谢产物合成的途径是稳定的,不同微生物物种的调节机制是可以弄清楚或基本弄清楚的,这就是对这种合成代谢流“治理”的基础。这种“治理”要用到遗传手段(遗传脱钩),并辅之以培养条件的优化(环境协调),以求达到初级代谢产物过量合成的目的。Ⅰ. 初级代谢产物过量合成的五字策略 在分析细胞机器的生产运行时,可把它看作是分步进行的过程。首先细胞外的营养物质从环境(培养基)跨膜进入细胞;然后,一般要经过向心、中心和离心 3 段连续的代谢途径的代谢,才能在胞内生成目的产物;最后,目的产物经过膜排出细胞,回到环境(发酵醪液)。P “载流途径”泛指承载代谢物流的代谢途径。“载流路径”泛指由不止一条载流途径按序首尾衔接而组成的代谢通道。 在发酵工程领域,“载流路径”作为发酵工程的专用术语,被定义为从主要有机原料到目的产物一组载流途径的有序组合。 典型的载流路径是指存在于代谢网络中的这样的代谢通道,即由多至五段承担不同代谢分工的、依次衔接的代谢途径组成的代谢通道。因此有载流路径的五段式之称。在这条载流路径上流动的代谢主流对应地也有五段,从而派生出代谢主流的五段式。载流途径的五段式和代谢主流的五段式统称为微生物代谢的五段式。 在此五段式的基础上,从不同的角度出发,提出能作为一个整体,用于设计育种以及发酵工艺控制的五字策略(简称工业发酵的五字策略):进、通、节、堵、出。“ 五字策略 ” 就是从合作的良好愿望出发,发挥人的主观能动作用,主动与微生物合作实现工业发酵效益的策略。这个策略来自初级代谢产物过量合成的研究,故实际上是初级代谢产物过量合成的五字策略。"工业发酵的“五字策略”PowerPoint 演示文稿E进:在育种和发酵控制方面都要促进细胞对营养物质的吸收。在发酵过程中,一般都希望菌体能迅速达到一定的浓度,因此,就必需选用生长旺盛的菌株,这类菌株吸收营养物质的能力往往也较强。同时要选择适当的容易吸收的培养基,和适宜的培养条件,以促进微生物的生长。为了消除底物在培养过程中的抑制作用,有的还采用流加的培养方法。为了促进微生物的生长和支持目的产物的合成,还要从代谢生理学的角度优化流加条件。 通:在育种方面,解除对途径中某些酶的反馈调节;在发酵控制方面,诱导这些酶的合成或激活这些酶,从而使来自各代谢物流(除碳架物流外还包括其他支持生物合成的物流)能够畅通地注入载流路径,汇入代谢主流,流向目的产物。特别是当发酵进入目的产物合成阶段后,必需确保载流路径畅通,代谢主流优势明显。   例如在谷氨酸的发酵过程中,作为谷氨酸合成原料的 NH4+ 的缺乏, 就可能使谷氨酸合成途径的中断,导致谷氨酸产量下降,大量α-酮戊二酸积累。当然,合成途径中的任何一个限制因子,都可能产生类似的情况。因此,在发酵过程中应尽可能的使通向载流路径的代谢途径畅通无阻,使不同的营养物质能顺利地注入代谢主流,最终流向目的产物。节:采用育种或发酵控制手段,节制载流路径上与目的产物的形成无关或关系不大的代谢支流,使碳架物质相对集中地流向目的产物。这里所谓的 “ 节制 ” 是指封闭或削弱以目的产物合成途径的起始底物(如下图中的A)或各中间产物(如下图中的B、C 和D)为起始底物的分支途径( 即下图中通向 K,G,I,J 的途径)。PowerPoint 演示文稿 经过千千万万年的物竞天择,微生物细胞已经成为自我保障的经济体系。细胞严格的调节机制不允许代谢中间产物的过量合成,要想实现目的产物的过量合成,就必需对其代谢进行导向(包括遗传根据和培养条件的导向),迫使代谢朝着过量合成目的产物方向进行。因此,不是目的产物生产所必需的途径,应做到不同程度的关闭,也就是“节”。堵:采用育种或发酵控制手段堵塞或削弱目的产物进一步代谢的途径(如上图中 F 以后的途径),包括目的产物参与的分解代谢和合成代谢。如果菌株生产目的产物的途径畅通,但同时目的产物进一步代谢(包括目的产物参与的分解代谢和合成代谢)的途径也畅通,或者随着生产的进行目的产物进一步的代谢加强,都会最终影响目的产物的溢出代谢,降低工业发酵的效益。  为了消除这种影响,就必须降低目的产物进一步代谢的酶的酶活力或酶量,以消除或削弱目的产物进一步的代谢。若不能对菌株进行有效的改造,也可以通过工艺手段来达到堵的目的。例如,采用及时将目的产物从发酵液的分离出来的办法,也可以间接地消除或削弱胞内目的产物的进一步代谢,甚至能拉动溢出代谢。=出:促进目的产物向胞外空间分泌。根据微生物细胞经济学原理(节约原理、适应原理、生命保障原理),如果细胞内某代谢中间产物浓度超过其阈值,必将受到细胞内相关代谢调节机制的干预。胞内目的产物的浓度过高,就会对合成目的产物的代谢过程施加负反馈调节,迫使其回到正常浓度。因此,及时将目的产物转移到细胞外,无论从化学平衡的角度,还是从生物学信息反馈的角度,都将有利于载流路径上代谢主流的流动。 ( 在育种和发酵控制方面加强“出”的办法很多,如调节细胞对目的产物的通透性,增加输送目的产物的载体蛋白的量,为目的产物的主动输送提供代谢能等,都能使在胞内生成的目的产物尽快的转移出细胞,降低其细胞中的浓度;为了进一步降低细胞中目的产物的浓度,还可能根据目的产物的理化性质,采用边发酵边分离目的产物的工艺,不断的将目的产物从反应器中转移出去,给代谢主流一个“拉动力”。Ⅱ.有关碳架物质跨膜问题的分析 氨基酸的发酵法生产,属于本课程定义的典型的工业发酵。以糖质原料发酵生产氨基酸的过程至少依次包括糖质进入细胞、经过代谢在胞内生成目的氨基酸和目的氨基酸被分泌至培养基 3 个阶段。此外,氨基酸的生产过程还包括细菌细胞的适度生长和生命活动的维持。 单考虑细胞内的碳架物质等的代谢是不够的,至少还必须考虑营养物质进入细胞及目的产物排出细胞的问题,也就是碳架物质通过细胞膜的问题。 有关碳架物质跨膜问题,必要时可做专题讲座。hⅢ.有关代谢途径的流量调节的分析 以天冬氨酸族氨基酸的合成代谢为例,来讨论代谢途径的流量调节。对细菌来说,天门冬氨酸族氨基酸包括Asp、Asn、Lys、The、Ile 和 Met 共6 种氨基酸( 或氨基酰胺),它们都是从 TCA 环中的前体代谢物草酰乙酸(OAA)开始合成的。 微生物细胞以葡萄糖为原料合成天冬氨酸族氨基酸时,碳架物质通过代谢网络的载流路径流向目的产物,其载流路径的上游部分从葡萄糖起到草酰乙酸止,草酰乙酸以后为下游部分。其上游部分属于中心途径,下游部分属于离心途径。因此天冬氨酸族氨基酸合成代谢流是跨代谢网络的中心板块和离心板块的。关于天冬氨酸族氨基酸的合成代谢路线(下游部分)及其调节方式已汇总在下面几张图中:PowerPoint 演示文稿PowerPoint 演示文稿PowerPoint 演示文稿 芳香族氨基酸的合成也有类似的情况。其载流路径的上游部分从葡萄糖起到磷酸烯醇式丙酮酸和4-磷酸赤藓糖止,以后为下游部分。其上游部分属于中心途径,下游部分属于离心途径。因此芳香族氨基酸合成代谢流是跨代谢网络的中心板块和离心板块的。关于芳香族氨基酸的合成代谢路线(下游部分)及其调节方式已汇总在下面几张图中:PowerPoint 演示文稿PowerPoint 演示文稿 下面是乳糖发酵短杆菌的赖氨酸高产菌株的设计育种示意图。绿色箭头连接成的路径为用葡萄糖为主要碳源时的理想载流路径。所谓 “ 理想载流路径 ” 是指为了获得高效益,人们期望代谢流流经的路径。PowerPoint 演示文稿A 生物合成途径同一性的一个例外:赖氨酸的生物合成途径在细菌是二氨基庚二酸途径,在真菌中是氨基己二酸途径。上图显示的是乳糖发酵短杆菌的合成赖氨酸的二氨基庚二酸途径,下图显示的是产黄青霉合成赖氨酸的氨基己二酸途径。PowerPoint 演示文稿*Ⅳ.应用“五字策略”的实施 “节” 的成功实施:若以黄色短杆菌为出发菌株,选育 L-Lys 高产菌种,可采用高丝氨酸营养缺陷型突变株。假设高丝氨酸营养缺陷是因为高丝氨酸脱氢酶( HD )缺损而造成的,从前面对应的图可找出造成 L-Lys高产的原因:①阻止碳架物质 Asp-β-SA 流向H-Ser;②解除 Thr、Lys对天门冬氨酸激酶( AK )的协同反馈抑制(因为细胞不能合成Thr);③省下原用于11e合成的PYR,以支持Lys合成。 从结构类似物的超敏性突变株中获得高产菌株是可行的,但必须注意这里 “ 结构类似物 ” 是指酶的底物的结构类似物,前述的“酶” 必须是与微生物的生长密切有关的酶。与微生物生长有密切关系的酶的底物的结构类似物,如果能竞争性地抑制这个酶的活性,就会直接影响微生物的生长,也就是这种微生物对这种结构类似表现出敏感性。这种敏感性的大小与酶水平的高低相关,敏感性小的菌株酶水平高,敏感性大的菌株酶水平低,特别敏感的(超敏的)的菌株酶水平最低。A 根据这个道理,有可能从这种酶的底物的结构类似物的超敏性突变株中,找到这种酶的低水平突变株。酶水平低,代谢流的通道就窄,就意味着“节”(节流)。 氟代丙酮酸( FP )是丙酮酸( PYR )的结构类似物。PYR是与生长密切相关的丙酮酸脱氢酶( PD ) 的底物,如果用葡萄糖为唯一碳源,PD 是唯一通向 AcCoA和TCA环的通道,若 PD 完全阻塞,微生物就无法通过TCA环和电子传递获得代谢能,有关的合成代谢也无法进行。 已用超敏性突变株的方法选育了乳糖发酵短杆菌的氟代丙酮酸超敏性突变株(FPs)。根据对FP超敏程度的不同,选得不同程度的PD缺损株( PD的不同程度的低水平株 ),从中找到了 PD水平低到恰到好处的突变株,既不过分削弱 PD 以提供足够的代谢能,又能使碳架物质较多地流向 OAA 以合成更多的Lys。从而实施了对PD通道的 “节”。  “通”的实施: 赖氨酸的结构类似物S-(β-aminoethyl)-L-Cys,简称 AEC,是最早用于氨基酸育种并取得明显效果的结构类似物。 在黄色短杆菌的亮氨酸缺陷型突变株 Leu-上再加上结构类似物抗性标记( AEC+Thr )r 后Lys产量大幅度上升,这是因为这个突变株已解除了Lys和Thr对天门冬氨酸激酶( AK )的协同反馈抑制,是个调节突变株。 关于“进”、“出”、“堵”的策略应用也都有成功的实例。 I⑵ 次级代谢产物的合成 人们一般把微生物营养性生长所需的代谢产物(直接或间接的参与细胞结构或组成)叫做初级代谢产物,把微生物在限制生长或没有生长的条件下形成的、对微生物的营养性生长并不必需的代谢产物叫做次级代谢产物。" 次级代谢产物种类很多,然而,它们的结构单位主要是莽草酸(SMA)、氨基酸 ( ar-AA 和 al-AA ) 和乙酸(AcCoA),也就是说它们可以从莽草酸、某些氨基酸和AcCoA等通过(有条件诱导的 )次级代谢途径来合成:PowerPoint 演示文稿Q 次级代谢产物的存在已引起人们特别注意。在典型的次级代谢中,次级代谢产物以混合物状态出现,即微生物可合成化学上相关联的一类次级代谢产物的混合物。显然,这是因为次级代谢所涉及的某些酶的特异性比初级代谢的酶的特异性要低得多。 次级代谢产物合成途径的型式也比初级代谢多得多。有人说次级代谢好像是有机物代谢演变的“操场”,这是有一定道理的。只要演变出来的次级代谢途径或其产物对途径的载体微生物没有害处,它们对应遗传物质就会随次级代谢途径的载体微生物的传代而得以保留或改善;反之,如果对途径的载体微生物有害,载体微生物本身就会被淘汰。B 换句话说,为次级代谢途径编码的DNA如果对细胞的任何一种组织水平,包括中间代谢、调节、输送、分化、形态发生等等 ,有好处,就会被保存下来,甚至得到扩展;如果没有用就有可能会被删除。抗生素的合成途径可能就是这样来的。 次级代谢一旦建立,就将服从寄主细胞固有的调节机制,并且有可能引伸出特殊的调节机制。在已建立的次级代谢体系中,曾发现直接或间接地作用于次级代谢的级联(cascade)调节效应。 抗生素是次级代谢产物中按功能而划定的复杂化合物类群。同一菌种可以合成结构相近的抗生素混合物,混合物中每种组分的比例决定于遗传和环境因素。 虽然次级代谢产物多种多样,但从生物化学的角度来看,大多数次级代谢产物都是从少数关键的中间代谢产物“ 组装 ” 的: PowerPoint 演示文稿 因此,用来汇入次级代谢前体库的代谢流对次级代谢是至关重要的。在这个意义上,研究初级代谢流及其调节,对次级代谢产物的合成代谢来说同样是很重要的。 次级代谢产物一般是在分批培养的分化期开始形成,即在限制生长或没有生长的条件下形成的。 次级代谢途径(酶)一般并不是组成型的,而是经诱导(或解除阻遏)而形成的。不能忽视这个事实: 放线菌与细菌相比较,是次级代谢产物的更好的生产者,放线菌有细胞分化的能力,而次级代谢往往与分化过程有关。 我们已讨论了某些关于代谢产物过量合成的生理学的一般规律。产品形成的生理学的基本道理是容易理解的。同时,我们已经注意到,在许多场合下,特别是关于次级代谢产物的生成,其生理学的知识仍是远远不够的。因此很有必要在此领域从事更多的研究,并要吸取在初级代谢生理学的类似研究中所得到的经验和教训。 已用的字体演示文稿设计模板嵌入 OLE 服务器幻灯片标题h_S**  !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      !"#$%&'()*+,-./0123456789:;<=>?@ABCDEFGHIJKLMNOPQRSTUVWXYZ[\]^_`abcdefghijklmnopqrstuvwxyz{|}~      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