Co-utilization of xylose and glucose to produce chemicals by microorganisms

doi:10.16085/j.issn.1000-6613.2022-0573

Abstract

Abstract:

Lignocellulosic biomass is abundant which has been proved to be a promising renewable resource in energy, chemical and pharmaceutical, etc. The efficient hydrolysis and utilization of lignocellulose are limited by the existence of hydrogen bond and covalent bond between its components, which require complicated pretreatment methods, such as acid, alkali, high temperature, organic solvent and enzymatic hydrolysis to overcome. The main component of lignocellulosic hydrolysates is the mixture of hexose (60%—70%, mainly glucose) and pentose (30%—40%, mainly xylose). Aiming at the utilization efficiency of xylose and glucose in the hydrolysates, this paper reviewed xylose and glucose co-fermentation in genetically engineered microorganisms for the production of alcohol, biolipids, γ-PGA and organic acids, and described the the research progress in reconstruction of metabolic pathways, regulations of gene level and fermentation technology optimization. Finally, the main characteristics of recent research, technical bottlenecks and future research directions in this field were summarized from the perspectives of strain screening, gene and metabolic engineering regulation, immobilization of cells, product treatment and fermentation process, in order to obtain more research methods and ideas.

Key words: xylose metabolism, glucose metabolism, fermentation, bioenergy, alcohol, organic acids

摘要：

木质纤维素生物质是储量丰富的可再生资源，在能源、化工及医药领域具有广阔的应用前景。木质纤维素各组分因氢键和共价键的存在而结合紧密，需经酸、碱、高温、有机溶剂等预处理后才能高效酶解利用，其水解产物的主要成分为己糖（60%~70%，葡萄糖为主）和戊糖（30%~40%，木糖为主）的混合物。本文主要针对水解液中木糖和葡萄糖的共利用效率相关问题，以基因工程改造微生物利用木糖和葡萄糖共发酵生产醇类、生物油脂、γ-聚谷氨酸及有机酸等生物基化学品为主线，从代谢途径重构、基因水平调控及发酵技术优化等方面综述了近年来的研究进展。最后，从菌株筛选、基因与代谢工程调控、细胞固定化、产物处理及发酵工艺等层面总结了该领域目前的研究特点、技术瓶颈和未来的研究方向与思路。

关键词: 木糖代谢, 葡萄糖代谢, 发酵, 生物能源, 醇, 有机酸

CLC Number:

Q819

WANG Chuandong, ZHANG Junqi, LIU Dingyuan, MA Yuanyuan, LI Feng, SONG Hao. Co-utilization of xylose and glucose to produce chemicals by microorganisms[J]. Chemical Industry and Engineering Progress, 2023, 42(1): 354-372.

王川东, 张君奇, 刘丁源, 马媛媛, 李锋, 宋浩. 微生物共利用木糖和葡萄糖生产化学品研究进展[J]. 化工进展, 2023, 42(1): 354-372.

Figures/Tables 5

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酶催化步骤	G^θ/kJ·mol^-1	∆G^θ/kJ·mol^-1
Step0：初始态	0	—
Step1：XDH	-19.20	-19.20
Step2：XLA	-40.40	-21.20
Step3：XAD	-82.30	-41.90
Step4：KDXD	-121.50	-39.20
Step5：KGSADH	-156.80	-35.30

酶催化步骤	G^θ/kJ·mol^-1	∆G^θ/kJ·mol^-1
Step0：初始态	0	—
Step1：XDH	-19.20	-19.20
Step2：XLA	-40.40	-21.20
Step3：XAD	-82.30	-41.90
Step4：KDXD	-121.50	-39.20
Step5：KGSADH	-156.80	-35.30

菌株	代谢途径	原料	技术方法	产物产量/g·L^-1	收率/g·g^-1	参考文献
T. cutaneum	EMP-TCA途径	玉米秸秆	选择培养基筛选，3L发酵罐BF	油脂	0.39	[5]
R. arrhizus	磷酸酮酶途径	木糖、葡萄糖	选择培养基筛选，SF	FA 46.78	0.47	[6]
C. tyrobutyricum	ED-XR途径	大豆壳、玉米纤维、小麦秸秆、水稻秸秆、甘蔗渣	表达异源XylT、XylA、XylB基因，1L搅拌槽生物反应器	丁酸42.60	0.36	[7]
E. coli	XR-XI途径	空棕榈果束纤维	适应性进化，基因突变筛选，表达戊糖代谢基因，混糖，微型发酵罐FBF	木糖醇4.80	0.99	[8]
K. marxianus	EMP途径	满江红	热酸水解、酶法糖化，半厌氧SF	乙醇26.80	0.43	[11]
C. lusitaniae	EMP途径	满江红	热酸水解、酶法糖化，半厌氧SF	乙醇23.20	0.37	[11]
S. stipitis	EMP途径	满江红	热酸水解、酶法糖化，半厌氧SF	乙醇18.20	0.29	[11]
P. tannophilus	XR途径	玉米芯	稀酸热水解、木碳粉解毒处理、滤纸过滤，Luedeking-Piret模型分析，深层发酵	木糖醇0.81	0.81	[13]
D. hansenii varhanseni	XR途径	玉米芯	稀酸热水解、木碳粉解毒处理、滤纸过滤，Luedeking-Piret模型分析，深层发酵	木糖醇0.79	0.79	[13]
C. guillermondii	XR途径	稻草	稀酸热水解、木碳粉解毒处理、滤纸过滤，Luedeking-Piret模型分析，深层发酵	木糖醇0.76	0.76	[13]
S. cerevisiaea C. utilis	EMP途径	芦苇秸秆	60目筛、热碱液处理，加纤维素酶，混菌有氧SF	乙醇9.81	0.10	[15]
B. subtilis	谷氨酸合成-WBG途径	木糖、葡萄糖	表达异源WBG途径基因，0.50L搅拌槽发酵罐BF	γ-PGA 5	0.26	[31]
CyanobacteriumSynechocystis	XI-EMP-TCA途径	木糖、葡萄糖、CO₂	糖原合成突变体∆glgC中表达木糖分解代谢基因XylA、XylB，CO₂固定，氮饥饿条件，SF	丙酮0.61、 AKG 0.42	0.76、 0.32	[45]
C. glutamicum	EMP-XI途径	木糖、葡萄糖	结合甘油和3-HP合成基因，筛选活性醛脱氢酶，iolT1和glk取代PEP依赖的PTS，整合araE和XylA/B，5L发酵罐FBF	3-HP 62.60	0.51	[46]
E. coli	XR-WBG途径	木糖、葡萄糖	表达异源合成基因Xdh、MdlC，敲除旁路基因XylA、YjhH和YagE，过表达Zwf，敲除MtfA，失活PntAB基因，SF	BT 7.23	0.55	[47]
K. pneumoniae	WBG途径	木糖、葡萄糖	表达Xdh、KivD及YjhG基因，敲除XylA基因，SF	BT 4.52	0.21	[48]
S. cerevisiae	WBG途径	木糖、葡萄糖	XylB、XylD、KdcA、Adh基因编码的异源四酶反应，整合KdcA基因至酵母基因组，发酵罐FBF	BT 6.60	0.57	[49]
K. marxianus	XR-EMP	木糖、葡萄糖	过表达异源XR、XD基因，敲除甘油、乙酸副产物基因，发酵罐FBF	乙醇51.43	0.35	[50]
K. marxianus	XR-WBG途径	木糖、葡萄糖	敲除Xyl1、Xyl2，过表达异源Xyl1基因，选择培养基筛选，无灭菌FBF	木糖醇 312.05	0.99	[50]
K. marxianus	XR途径	木糖、葡萄糖	过表达异源Xyl1基因，敲除木糖醇代谢基因，重构葡萄糖代谢，FBF	木糖醇203.57	0.99	[51]
Y. lipolytica	EMP-XR途径	木糖、葡萄糖	表达外源XDH、XR和内源XK、GPD1和DGA2，敲除POX1-6和TGL4基因，5L发酵罐FBF	油脂22.50、柠檬酸67.20	0.06、 0.18	[52]
T. laichii	EMP-TCA	木糖、葡萄糖	选择培养基筛选，FBF	油脂	0.83	[53]
R. toruloides	EMP-TCA	木糖、葡萄糖	选择培养基筛选，FBF	油脂	0.65	[53]
L. starkeyi	从头合成途径	玉米芯	稀酸处理，过碱化和活性炭吸附解毒，SF	油脂	0.47	[54]
C. humicola	从头合成途径	玉米秸秆	高温、高压、碱水解预处理，SF	油脂	0.44	[54]
B. amyloliquefaciens	XI-不依赖谷氨酸的谷氨酸合成途径	玉米秸秆、豆粕	50L、150L曝气式连续搅拌固态生物反应器CF	γ-PGA 117、102*	—	[55]
R. arrhizus	磷酸酮酶途径	木糖、葡萄糖	定向进化、SF	FA 28.48	0.46	[56]
R. delemar	EMP-TCA途径	葡萄糖、玉米浆	粒径0.55mm球团固定、搅拌槽发酵罐BF	FA 39.56	0.40	[57]
R. oryzae	LA发酵	木糖、葡萄糖	诱变筛选，SF	乳酸119.22	0.80	[58]
B. coagulans	LA发酵	玉米芯残渣	SSF、FBF	乳酸68	0.85	[59]
B. coagulans	LA发酵	麦秸	稀硫酸预处理，7.50L发酵罐SSF	乳酸52.20	0.87	[60]
B. coagulans	LA发酵	木薯、高粱粉	补加α-淀粉酶、糖化酶10L发酵罐SSF	乳酸68.72	0.99	[61]
C. tyrobutyricum	ED-XR途径	木糖、葡萄糖	表达异源XylT、XylA、XylB基因，补加紫精，1L发酵罐BF	丁酸46.40	0.43	[62]
K. marxianus	XR-EMP途径	玉米芯	稀酸处理、微曝气两段式，5L发酵罐SSF、FBF	乙醇24.20	0.82	[63]
K. marxianus	XR-EMP途径	玉米芯	稀酸预处理，5L发酵罐微曝、两段式SSF、FBF	木糖醇52	0.41	[63]

菌株	代谢途径	原料	技术方法	产物产量/g·L^-1	收率/g·g^-1	参考文献
T. cutaneum	EMP-TCA途径	玉米秸秆	选择培养基筛选，3L发酵罐BF	油脂	0.39	[5]
R. arrhizus	磷酸酮酶途径	木糖、葡萄糖	选择培养基筛选，SF	FA 46.78	0.47	[6]
C. tyrobutyricum	ED-XR途径	大豆壳、玉米纤维、小麦秸秆、水稻秸秆、甘蔗渣	表达异源XylT、XylA、XylB基因，1L搅拌槽生物反应器	丁酸42.60	0.36	[7]
E. coli	XR-XI途径	空棕榈果束纤维	适应性进化，基因突变筛选，表达戊糖代谢基因，混糖，微型发酵罐FBF	木糖醇4.80	0.99	[8]
K. marxianus	EMP途径	满江红	热酸水解、酶法糖化，半厌氧SF	乙醇26.80	0.43	[11]
C. lusitaniae	EMP途径	满江红	热酸水解、酶法糖化，半厌氧SF	乙醇23.20	0.37	[11]
S. stipitis	EMP途径	满江红	热酸水解、酶法糖化，半厌氧SF	乙醇18.20	0.29	[11]
P. tannophilus	XR途径	玉米芯	稀酸热水解、木碳粉解毒处理、滤纸过滤，Luedeking-Piret模型分析，深层发酵	木糖醇0.81	0.81	[13]
D. hansenii varhanseni	XR途径	玉米芯	稀酸热水解、木碳粉解毒处理、滤纸过滤，Luedeking-Piret模型分析，深层发酵	木糖醇0.79	0.79	[13]
C. guillermondii	XR途径	稻草	稀酸热水解、木碳粉解毒处理、滤纸过滤，Luedeking-Piret模型分析，深层发酵	木糖醇0.76	0.76	[13]
S. cerevisiaea C. utilis	EMP途径	芦苇秸秆	60目筛、热碱液处理，加纤维素酶，混菌有氧SF	乙醇9.81	0.10	[15]
B. subtilis	谷氨酸合成-WBG途径	木糖、葡萄糖	表达异源WBG途径基因，0.50L搅拌槽发酵罐BF	γ-PGA 5	0.26	[31]
CyanobacteriumSynechocystis	XI-EMP-TCA途径	木糖、葡萄糖、CO₂	糖原合成突变体∆glgC中表达木糖分解代谢基因XylA、XylB，CO₂固定，氮饥饿条件，SF	丙酮0.61、 AKG 0.42	0.76、 0.32	[45]
C. glutamicum	EMP-XI途径	木糖、葡萄糖	结合甘油和3-HP合成基因，筛选活性醛脱氢酶，iolT1和glk取代PEP依赖的PTS，整合araE和XylA/B，5L发酵罐FBF	3-HP 62.60	0.51	[46]
E. coli	XR-WBG途径	木糖、葡萄糖	表达异源合成基因Xdh、MdlC，敲除旁路基因XylA、YjhH和YagE，过表达Zwf，敲除MtfA，失活PntAB基因，SF	BT 7.23	0.55	[47]
K. pneumoniae	WBG途径	木糖、葡萄糖	表达Xdh、KivD及YjhG基因，敲除XylA基因，SF	BT 4.52	0.21	[48]
S. cerevisiae	WBG途径	木糖、葡萄糖	XylB、XylD、KdcA、Adh基因编码的异源四酶反应，整合KdcA基因至酵母基因组，发酵罐FBF	BT 6.60	0.57	[49]
K. marxianus	XR-EMP	木糖、葡萄糖	过表达异源XR、XD基因，敲除甘油、乙酸副产物基因，发酵罐FBF	乙醇51.43	0.35	[50]
K. marxianus	XR-WBG途径	木糖、葡萄糖	敲除Xyl1、Xyl2，过表达异源Xyl1基因，选择培养基筛选，无灭菌FBF	木糖醇 312.05	0.99	[50]
K. marxianus	XR途径	木糖、葡萄糖	过表达异源Xyl1基因，敲除木糖醇代谢基因，重构葡萄糖代谢，FBF	木糖醇203.57	0.99	[51]
Y. lipolytica	EMP-XR途径	木糖、葡萄糖	表达外源XDH、XR和内源XK、GPD1和DGA2，敲除POX1-6和TGL4基因，5L发酵罐FBF	油脂22.50、柠檬酸67.20	0.06、 0.18	[52]
T. laichii	EMP-TCA	木糖、葡萄糖	选择培养基筛选，FBF	油脂	0.83	[53]
R. toruloides	EMP-TCA	木糖、葡萄糖	选择培养基筛选，FBF	油脂	0.65	[53]
L. starkeyi	从头合成途径	玉米芯	稀酸处理，过碱化和活性炭吸附解毒，SF	油脂	0.47	[54]
C. humicola	从头合成途径	玉米秸秆	高温、高压、碱水解预处理，SF	油脂	0.44	[54]
B. amyloliquefaciens	XI-不依赖谷氨酸的谷氨酸合成途径	玉米秸秆、豆粕	50L、150L曝气式连续搅拌固态生物反应器CF	γ-PGA 117、102*	—	[55]
R. arrhizus	磷酸酮酶途径	木糖、葡萄糖	定向进化、SF	FA 28.48	0.46	[56]
R. delemar	EMP-TCA途径	葡萄糖、玉米浆	粒径0.55mm球团固定、搅拌槽发酵罐BF	FA 39.56	0.40	[57]
R. oryzae	LA发酵	木糖、葡萄糖	诱变筛选，SF	乳酸119.22	0.80	[58]
B. coagulans	LA发酵	玉米芯残渣	SSF、FBF	乳酸68	0.85	[59]
B. coagulans	LA发酵	麦秸	稀硫酸预处理，7.50L发酵罐SSF	乳酸52.20	0.87	[60]
B. coagulans	LA发酵	木薯、高粱粉	补加α-淀粉酶、糖化酶10L发酵罐SSF	乳酸68.72	0.99	[61]
C. tyrobutyricum	ED-XR途径	木糖、葡萄糖	表达异源XylT、XylA、XylB基因，补加紫精，1L发酵罐BF	丁酸46.40	0.43	[62]
K. marxianus	XR-EMP途径	玉米芯	稀酸处理、微曝气两段式，5L发酵罐SSF、FBF	乙醇24.20	0.82	[63]
K. marxianus	XR-EMP途径	玉米芯	稀酸预处理，5L发酵罐微曝、两段式SSF、FBF	木糖醇52	0.41	[63]

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