微生物学报

材料 Material	单体 Monomer	常见聚合方式 Common polymerization methods	物理性能 Physical performance	参考文献References
Polycaprolactone (PCL)	己内酯 ε-caprolactone	Open-loop polymerization method	Exhibits depressed thermal transitions with a melting point (T_m=60 ℃) and glass transition temperature (T_g=-60 ℃), demonstrating exceptional processability	[14]
Polylactic acid (PLA)	乳酸(或丙交酯) Lactic acid	Polymerization, open-loop polymerization method, enzymatic polymerization	Distinctive thermal stability manifested through elevated T_m (170 ℃) and T_g (60 ℃) exceeding conventional aliphatic polyester systems	[15]
Polyhydroxyalkanoates(PHA)	多种羟基脂肪酸 Hydroxyalkanoic acid	-	Presents a T_mof 145 ℃ with adjustable physicochemical properties through monomer compositional engineering and stoichiometric optimization	[16]
Poly(β- hydroxybutyrate) (PHB)	3-羟基丁酸 3-hydroxybutyric acid	Displays substantially elevated T_m(180 ℃) concomitant with superior mechanical rigidity and pronounced brittle fracture behavior	[17-18]
Poly(3-hydroxybutyrate-3-hydroxyvalerate (PHBV)	3-羟基丁酸 3-hydroxybutyric acid 3-羟基戊酸 3-hydroxyvaleric acid	Enables precise property tailoring via controlled 3-hydroxyvalerate content modulation in copolymer architecture	[19]
Poly(butylene adipate-co-terephthalate (PBAT)	己二酸 Adipic acid 对苯二甲酸Terephthalic acid 1,4-丁二醇 1,4-butanediol	Melt polycondensation	Thermal profile comprises: T_m= 110-130 ℃, T_g=-30 ℃, crystallization temperature (T_c) 110 ℃, and crystallinity degree (X_c) 30%	[20-21]
Poly propylene carbonate (PPC)	二氧化碳Carbon dioxide 环氧丙烷Propylene oxide	Coordination copolymerization	Features markedly elevated phase-transition temperatures with T_m=160-170 ℃ and T_g=30-41 ℃	[22]

材料 Material	单体 Monomer	常见聚合方式 Common polymerization methods	物理性能 Physical performance	参考文献References
Polycaprolactone (PCL)	己内酯 ε-caprolactone	Open-loop polymerization method	Exhibits depressed thermal transitions with a melting point (T_m=60 ℃) and glass transition temperature (T_g=-60 ℃), demonstrating exceptional processability	[14]
Polylactic acid (PLA)	乳酸(或丙交酯) Lactic acid	Polymerization, open-loop polymerization method, enzymatic polymerization	Distinctive thermal stability manifested through elevated T_m (170 ℃) and T_g (60 ℃) exceeding conventional aliphatic polyester systems	[15]
Polyhydroxyalkanoates(PHA)	多种羟基脂肪酸 Hydroxyalkanoic acid	-	Presents a T_mof 145 ℃ with adjustable physicochemical properties through monomer compositional engineering and stoichiometric optimization	[16]
Poly(β- hydroxybutyrate) (PHB)	3-羟基丁酸 3-hydroxybutyric acid	Displays substantially elevated T_m(180 ℃) concomitant with superior mechanical rigidity and pronounced brittle fracture behavior	[17-18]
Poly(3-hydroxybutyrate-3-hydroxyvalerate (PHBV)	3-羟基丁酸 3-hydroxybutyric acid 3-羟基戊酸 3-hydroxyvaleric acid	Enables precise property tailoring via controlled 3-hydroxyvalerate content modulation in copolymer architecture	[19]
Poly(butylene adipate-co-terephthalate (PBAT)	己二酸 Adipic acid 对苯二甲酸Terephthalic acid 1,4-丁二醇 1,4-butanediol	Melt polycondensation	Thermal profile comprises: T_m= 110-130 ℃, T_g=-30 ℃, crystallization temperature (T_c) 110 ℃, and crystallinity degree (X_c) 30%	[20-21]
Poly propylene carbonate (PPC)	二氧化碳Carbon dioxide 环氧丙烷Propylene oxide	Coordination copolymerization	Features markedly elevated phase-transition temperatures with T_m=160-170 ℃ and T_g=30-41 ℃	[22]

材料 Material	生物可降解地膜类型 Types of biodegradable mulch film	菌株 Strain	菌株来源 Source	降解周期 Degradation period (d)	降解率 Degradation rate	参考文献References
PCL	PCL mulch film	Pseudomonas sp. DS0901	Activated sludge	3	100%	[32]
PLA	PLA mulch film	Pseudomonas geniculata WS3	Soil and wastewater sludge	20	Around 45%	[42]
PLA mulch film	Stenotrophomonas pavanii CH1	/	20	10%	[42]
PLA mulch film	Bacillus pumilus B12	Soil	2	/	[76]
PLA powder/tray	Actinomadura keratinilytica T16-1	/	3	99%/32%	[77]
PLA mulch film	Nocardiopsis prasina	Ocean	60	(1.27±0.19)%	[78]
PLA mulch film	Priestia flexa PMPHB7	Soil	21	73%	[79]
PLA mulch film	Bacillus safensis PLA1006	Soil	30	8%	[80]
PLA mulch film	Tritirachium album	/	4	78%	[81]
PHA	PHA mulch film	/	Soil	80	75%	[48]
PHB	PHB mulch film	Acinetobacter junii BP25	Wastewater	77	83%	[82]
PHBV	PHBV	Alcaligenes spp. Pseudomonas spp.	Soil	365	13%	[83]
PBAT	PBAT mulch film	Enterobacter hormaechei WX-2	Soil	60	(20.8±2.2)%	[60]
PBAT mulch film (PF, AF, PPSF)	Thermobifida fusca FXJ-1	Compost samples	9	(82.87±1.01)%, (87.83±2.00)%, (52.53±0.54)%	[61]
PPC	PPC mulch film	Bacillus subtilis J16	Soil	30	9.95%	[73]

材料 Material	生物可降解地膜类型 Types of biodegradable mulch film	菌株 Strain	菌株来源 Source	降解周期 Degradation period (d)	降解率 Degradation rate	参考文献References
PCL	PCL mulch film	Pseudomonas sp. DS0901	Activated sludge	3	100%	[32]
PLA	PLA mulch film	Pseudomonas geniculata WS3	Soil and wastewater sludge	20	Around 45%	[42]
PLA mulch film	Stenotrophomonas pavanii CH1	/	20	10%	[42]
PLA mulch film	Bacillus pumilus B12	Soil	2	/	[76]
PLA powder/tray	Actinomadura keratinilytica T16-1	/	3	99%/32%	[77]
PLA mulch film	Nocardiopsis prasina	Ocean	60	(1.27±0.19)%	[78]
PLA mulch film	Priestia flexa PMPHB7	Soil	21	73%	[79]
PLA mulch film	Bacillus safensis PLA1006	Soil	30	8%	[80]
PLA mulch film	Tritirachium album	/	4	78%	[81]
PHA	PHA mulch film	/	Soil	80	75%	[48]
PHB	PHB mulch film	Acinetobacter junii BP25	Wastewater	77	83%	[82]
PHBV	PHBV	Alcaligenes spp. Pseudomonas spp.	Soil	365	13%	[83]
PBAT	PBAT mulch film	Enterobacter hormaechei WX-2	Soil	60	(20.8±2.2)%	[60]
PBAT mulch film (PF, AF, PPSF)	Thermobifida fusca FXJ-1	Compost samples	9	(82.87±1.01)%, (87.83±2.00)%, (52.53±0.54)%	[61]
PPC	PPC mulch film	Bacillus subtilis J16	Soil	30	9.95%	[73]

微生物降解聚酯和聚碳酸酯类地膜的研究进展

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王玉 ¹^,² , 王琰 ²^,^* , 聂红云 ¹^,³^,^* , 姚建民 ⁴ , 李瑞珍 ⁴ , 万一 ²

微生物学报 | 综述 2025,65(6): 2365-2381

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微生物学报 | 综述 2025, 65(6): 2365-2381

微生物降解聚酯和聚碳酸酯类地膜的研究进展

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王玉¹^,², 王琰²^,^*, 聂红云¹^,³^,^*, 姚建民⁴, 李瑞珍⁴, 万一²

作者信息

^1.西安建筑科技大学环境与市政工程学院，陕西西安

^2.陕西省微生物研究所，陕西西安

^3.陕西省秦岭生态安全重点实验室，陕西西安

^4.山西农业大学农业经济管理学院，山西太原

Advances in microbial degradation of polyester and polycarbonate-based mulch films

Yu WANG¹^,², Yan WANG²^,^*, Hongyun NIE¹^,³^,^*, Jianmin YAO⁴, Ruizhen LI⁴, Yi WAN²

Affiliations

^1.School of Environmental and Municipal Engineering, Xi’an University of Architecture and Technology, Xi’an, Shaanxi, China

^2.Shaanxi Institute of Microbiology, Xi’an, Shaanxi, China

^3.Shaanxi Key Laboratory of Qinling Ecological Security, Xi’an, Shaanxi, China

^4.College of Agricultural Economics and Management, Shanxi Agricultural University, Taiyuan, Shanxi, China

出版时间: 2025-06-04 doi: 10.13343/j.cnki.wsxb.20250044

文章导航

摘要

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生物可降解地膜(biodegradable mulch films, BDMs)以其广阔的应用前景和生态友好特性正逐步取代传统地膜，被视为解决“白色污染”问题最具潜力的途径。近年来，我国在生物可降解地膜的生产技术领域取得了显著成就，为其规模化生产和广泛应用奠定了坚实基础。尽管前景广阔，生物可降解地膜在降解过程中的复杂性、可控性以及其对生态环境的潜在影响仍是必须高度关注的重点。基于此，本文综合分析了5种极具应用前景的聚酯类和聚碳酸酯类生物可降解地膜，深入探讨了这些地膜的主要降解微生物及其降解机制，并对其土壤生态影响的研究现状进行了总结。本文为挖掘高效降解微生物资源、明确降解过程中的关键限速步骤、加强长周期生态效应研究提供了理论参考，为生物可降解地膜的大规模安全应用提供了新的解决思路和解决方案。

关键词

生物降解 / 聚酯 / 聚碳酸酯 / 酶 / 土壤生态

Abstract

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Biodegradable mulch films (BDMs), distinguished by their extensive application potential and ecological friendliness, are progressively supplanting traditional mulch film and considered as a highly promising approach to address “white pollution”. China has witnessed notable advancements in the production technology of BDMs in recent years, establishing a strong foundation for their large-scale manufacturing and widespread application. Despite the great prospects of BDMs, the complexity and controllability of their degradation process, alongside their potential impacts on the eco-environment, remain highly concerned. This paper comprehensively analyzes five promising polyester and polycarbonate-based BDMs and delves into the primary degrading microorganisms and their degradation mechanisms. Furthermore, this paper summarizes the current research regarding the impacts of BDMs on the soil environment. This review aims to lay a theoretical foundation for discovering efficient microbial degraders, pinpointing key rate-limiting steps in degradation, and enhancing long-term ecological effect studies, thus providing new perspectives and solutions for the large-scale and safe utilization of BDMs.

Key words

biodegradation / polyester / polycarbonate / enzyme / soil ecology

引用本文

王玉, 王琰, 聂红云, 姚建民, 李瑞珍, 万一. 微生物降解聚酯和聚碳酸酯类地膜的研究进展. 微生物学报, 2025 , 65 (6) : 2365 -2381 . DOI: 10.13343/j.cnki.wsxb.20250044

Yu WANG, Yan WANG, Hongyun NIE, Jianmin YAO, Ruizhen LI, Yi WAN. Advances in microbial degradation of polyester and polycarbonate-based mulch films[J]. Acta Microbiologica Sinica, 2025 , 65 (6) : 2365 -2381 . DOI: 10.13343/j.cnki.wsxb.20250044

正文

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地膜(mulch films)作为一种在农业生产中广泛应用的地面覆盖材料，自20世纪70年代末引入我国以来，凭借其增温、保墒、保肥、抑制杂草生长及改善土壤理化性质等多重功能，极大地推动了农业生产的发展^[1-2]。进入21世纪以来，地膜覆盖技术在我国历经不断探索与实践，已逐步迈向成熟阶段^[3]。然而，随着地膜覆盖技术的广泛应用，其在提升农业生产力的同时，也引发了日益严峻的残膜污染问题。传统地膜以聚乙烯(polyethylene, PE)、聚氯乙烯(polyvinyl chloride, PVC)等合成材料为基材，这些材料因高分子链结构的化学惰性，在自然环境中表现出极高的稳定性和难降解特性^[4]。在连续耕作模式下，土壤中地膜残留量持续累积，不仅严重破坏了土壤结构，降低了土壤透气性和孔隙度，还对作物的生长发育构成了极大威胁，包括降低出苗率、阻碍根系扩展及营养吸收等^[5-6]。尽管理论上可以通过回收地膜来解决这一问题，但在实际操作中，由于地膜材质的特性、回收成本及渠道等多重因素的影响，回收工作难以有效实施。作为农业大国，我国的地膜产量和使用量均居世界前列^[7]，由此引发的残膜污染问题愈发严峻，亟待解决。

生物可降解地膜(biodegradable mulch films, BDMs)凭借其环境友好性、可自然降解性，被视为解决地膜污染问题最有效的途径之一^[8]。近年来，我国在生物可降解地膜生产技术方面取得了显著进展，地膜性能得到大幅提升，使用成本逐步降低。例如，我国自主研发的0.006 mm聚碳酸亚丙酯(polypropylene carbonate, PPC)基生物降解地膜成功攻克了纵向撕裂强度小和耐候期短的共性技术难题，实现了万吨级PPC原料的生产，地膜中70%的PPC含量和0.006 mm的薄度极大降低了地膜成本与每亩用量^[9]，为其规模化生产及应用奠定了坚实基础。然而，生物可降解地膜在降解过程中的复杂性仍需高度关注，其逐级分解过程中可能形成诸如酸类、醚类及酯类，甚至微塑料等中间产物；同时，生物可降解地膜存在降解的不可预期性，可能出现作物需求期内即发生降解，以及作物收获后仍大面积残留的情况，从而影响下茬作物的耕种^[10]。这些逐级分解过程中可能形成的中间产物对土壤生态的影响，以及降解速率与作物生长周期的匹配性问题，均是当前研究的重点和难点。

基于此，本文综述了农业领域中广泛应用或具有广阔应用前景的5种聚酯类和聚碳酸酯类生物可降解地膜，探讨了这些地膜在农业环境中的主要降解微生物、降解酶、降解机制以及对土壤生态的影响。从环境中筛选出具有更强生物降解能力的微生物资源，深入挖掘潜在的生物降解机制，可为生物可降解地膜在农业生产中的安全应用与推广提供坚实的理论支撑，为解决地膜污染问题、实现可生物降解地膜的可控性提供新的思路和解决方案。

1 生物可降解地膜的种类

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生物可降解地膜是指一类在自然环境条件(包括但不限于土壤的湿度、温度、微生物群落等复杂生态条件)以及特定人工模拟环境条件(例如堆肥过程中适宜的高温、高湿和丰富微生物种群的环境，或是厌氧消化所营造的无氧且富含特定厌氧微生物的环境)下，发生自然水解反应致使覆盖膜聚合物化学键断裂分解，通过真菌或细菌分泌的特异性酶引起降解，直至最终完全降解为二氧化碳(CO₂)和水(H₂O)^[11-12]。目前，农业生产中广泛应用的生物可降解地膜以聚酯类和聚碳酸酯类为主^[13]，包括聚己内酯(polycaprolactone, PCL)、聚乳酸(polylactic acid, PLA)、聚羟基脂肪酸酯(polyhydroxyalkanoates, PHA)、聚己二酸/对苯二甲酸丁二酯[poly(butylene adipate-co-terephthalate), PBAT]和聚碳酸亚丙酯(polypropylene carbonate, PPC)。表1总结了这些生物可降解地膜的基本特征及性能。

2 作用于BDMs的微生物及酶

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目前广泛应用的生物可降解地膜均具有潜在的可水解键，易被微生物分泌的脂肪酶、酯酶、角质酶等解聚酶水解，并转变为低聚物、二聚体或单体，再进一步经同化与矿化作用转化为CO₂和H₂O^[12,23]。在已发现的参与降解BDMs的微生物类群中，真菌在种类数量上占优势。哈茨木霉、里氏木霉、链格孢霉、雅致放射毛霉、尖孢镰孢菌、黄曲霉、米曲霉、念珠菌等均是常见的具有降解能力的真菌菌株；在细菌方面，芽孢杆菌、大肠埃希氏菌、黄杆菌、假单胞菌、诺卡氏菌、克雷伯氏菌、红球菌、小单孢菌、放线菌等是主要的降解类群^[24-25]。本文将依据BDMs种类对其降解微生物以及所分泌的解聚酶进行介绍。

2.1 作用于PCL地膜的降解微生物及酶

PCL是通过ε-己内酯单体的开环聚合反应合成的，其分子链中的酯键能在多种微生物的作用下发生水解，从而实现生物降解^[23]。通过系统性分析发现，当前研究在PCL生物降解的微生物作用机制与酶促反应动力学领域取得了3项关键进展^[26-28]。

(1) 在环境响应特异性机制研究方面，Al Hosni等^[26]通过构建温度阶梯模型，揭示了微生物群落的温度适应性分化规律：在50 ℃堆肥体系中，成功分离出以绿脓杆菌为主的嗜热菌群(占比82.6%)，其最适生长温度[T_m=(48±1) ℃]与堆肥高温期显著相关；而在25 ℃的中温条件下，镰孢菌属成为优势菌群(相对丰度76.3%)，并首次证实新赤壳属(Neocosmospora sp.)菌株对PCL地膜的降解效率可达0.32 mg/(cm²⋅d)，相较对照组提升4.7倍。值得注意的是，高温菌群与中温菌群在降解功能上呈现出互补特征。这种由温度驱动的菌群演替规律，为设计具备梯度特性的堆肥系统提供了坚实的生物学理论依据。

(2) 对极端环境中微生物资源的深入开发，为PCL的降解研究开拓了全新的视角。Soulenthone等^[27]从海水中分离出一株嗜盐假单胞菌(Halopseudomonas sp.) MFKK-1，在海水盐度条件下能够维持(198.2±31.7) µg/(cm²⋅d)的降解速率。这一重要发现，不仅证实了海洋微生物在PCL降解过程中的关键作用，还暗示了海洋与陆地微生物在降解机制方面可能存在趋同进化的特征。此外，这些菌株所具备的高盐适应性是否归因于其酯酶的特殊修饰，值得进一步探究。

(3) 研究表明大多聚酯类材料的降解是由微生物胞外酯酶介导的^[28]。目前，已经从细菌或真菌中分离纯化出多种PCL解聚酶^[29]，其中最为关键的突破在于对这些酶解机制在分子层面的精准解析。Ma等^[30]对酶解机制展开了深入研究，发现PCL薄膜在南极假丝酵母脂肪酶作用下，其降解过程可清晰划分为快速、缓慢2个阶段，且PCL薄膜的分子量并不会对脂肪酶性能产生影响，却会显著影响降解后薄膜的孔结构。Feng等^[31]通过宏基因组筛选和多尺度模拟手段，揭示了PCL酶解的双步解聚机制，发现降解酶MGS0156能够通过疏水界面锚定在PCL表面，并通过构象转换驱动“表层剥离-纵深渗透”的降解路径。这一发现不仅构建了PCL酶解动力学的定量模型，还明确了C-O键断裂作为速率决定步骤的重要性，为酶工程的改造提供了清晰且明确的靶点，有望通过定向进化提升降解酶的催化效率。于丹等^[32]发现假单胞菌(Pseudomonas sp.) DS0901分泌的PCL解聚酶在37 ℃、130 r/min的摇床振荡培养条件下，72 h内可将PCL完全降解，并发现其立体降解模式与MGS0156作用机制存在耦合现象，这暗示着不同的解聚酶或许遵循着相同的降解规律。这一重要发现为建立具有普适性的聚酯降解理论模型奠定了坚实的基础。

然而，当前的研究大多集中在单一菌株或酶的降解性能上，而自然环境中微生物群落的代谢网络协同机制尚不明确。Tseng等^[33]从堆肥体系中分离出能够同时降解PCL、聚3-羟基丁酸酯[poly(β-hydroxybutyrate), PHB]等聚酯材料的降解菌(分属于小双孢菌属、高温放线菌属、马杜拉放线菌属、链霉菌属及木聚糖单胞菌属等)，为微生物群落水平的降解机制提供了研究模型。虽然PCL的生物降解研究在环境响应特异性机制、酶促反应动力学以及微生物群落协同机制等方面取得了重要进展，但是菌株是否通过代谢分工实现降解强化，微生物群体代谢是否参与降解酶的调控表达等许多问题仍有待深入探讨和研究。对这些问题的解答将有助于推动合成微生物群落的构建工作，实现降解效能的可控性。

2.2 作用于PLA地膜的降解微生物及酶

在降解PLA地膜的研究中，已发现了一系列关键的微生物及其分解酶。降解微生物主要包括5个细菌属：糖丝菌属、假诺卡氏菌属、链霉菌属、芽孢杆菌属和假单胞菌属，其中放线菌占据主导地位。此外，还包括真菌(如根霉、梗霉)和酵母菌(如隐球酵母)。对PLA地膜有突出降解能力的微生物主要集中于放线菌门的拟无枝酸菌属。Ikura等^[34]分离的拟无枝酸菌(Amycolatopsis sp.) 3118菌株，在12 d内对PLA的降解率可达到90%以上；Pranamuda等^[35]通过PLA乳化矿盐平板从土壤筛选出一株Amycolatopsis sp. HT-32，该菌株经过14 d的摇瓶培养可使60%的PLA薄膜发生降解，并且该降解过程被推测为酶促过程；Nakamura等^[36]从300份土壤样品中分离筛选到的Amycolatopsis sp. K104-1菌株，能以PLA为唯一碳源生长，并首次纯化出新型PLA降解酶——纤溶丝氨酸酶PLD，证实了PLA的酶解过程。Li等^[37]从东方拟无枝酸菌亚种(Amycolatopsis orientalis subsp. orientalis)中同时纯化出3种细胞外丝氨酸蛋白酶PLAase I、PLAase II、PLAase III，且3种酶均具良好的降解效果，进一步表明PLA的降解酶可能作为一种蛋白酶。

Williams等^[38]对分离自丝状真菌(Engyodontium album, 旧称Tritirachium album)的蛋白酶K进行了PLA生物降解性测试。Oda等^[39]对56种市售蛋白酶进行了PLA降解活性检测，发现嗜酸蛋白酶几乎对PLA无影响，中性蛋白酶仅有很低的降解活性，而部分嗜碱蛋白酶具有显著的聚乳酸降解活性；这些嗜碱蛋白酶大多来自芽孢杆菌属，即使在中性和碱性条件下也表现出催化活性，这可能与酶的空间结构和催化机制有关。Matsuda等^[40]纯化了一种蛋白酶，可使PLA薄膜在48 h内粉末化，降解率高达90%。此外，来自戴尔根霉(Rhizopus delemer)和解淀粉类芽孢杆菌(Paenibacillus amylolyticus) TB-13的脂肪酶，来自史氏芽孢杆菌(Bacillus smithii)的酯酶以及来自食酸代尔夫特菌(Delftia acidovorans) TB-35菌株的聚酯聚氨酯降解酶均被报道对PLA有降解能力^[41-43]。Akutsu-Shigeno等^[44]对来自P. amylolyticus TB-13的PLA降解基因进行克隆，并在大肠埃希氏菌(Escherichia coli)中进行异源表达和纯化，纯化获得的重组PLA解聚酶PlaA表现出对多种可降解聚酯(如聚丁二酸丁二醇酯、聚丁二酸乙二醇酯和聚己内酯等)的降解活性，其在90 min内可完全降解分子量为5 000 g/mol的PLA，并且降解效率远高于蛋白酶K。重组脂肪酶展现出的高降解性，不仅为聚酯类废弃物的绿色处理提供了新的酶学途径，也证实广泛存在于自然环境中的产脂肪酶微生物是PLA的潜在降解类群。

此外，Watanabe等^[45]从稻壳中分离出隐球菌(Cryptococcus sp.) GB-1，它可产生可生物降解塑料降解酶，采用紫外线诱导获得突变体GB-1-DMC1，其在葡萄糖/木糖双碳源体系中PLA降解酶活性较原始菌株提升3.1倍，这一现象提示碳源代谢调控可能通过表观遗传修饰影响酶蛋白表达水平。值得注意的是，蛋白酶底物特异性研究表明^[46]，蛋白酶对l-乳酸有较高的降解活性，而对d-乳酸表现出较高的惰性，并且对聚乳酸的结晶度具有很高的敏感性。这种差异可能与酶的活性中心与PLA构象的匹配模式有关。这一发现为进一步优化酶的立体选择催化位点提供了新思路。尽管在PLA降解方面已经取得了较多进展，但仍有许多问题亟待解决。例如，如何通过定向进化策略优化酶的立体选择催化位点，如何借助代谢网络重构技术提升工程菌株的酶蛋白表达量及分泌效率等。这些问题的解答将有助于更深入地理解PLA的微生物降解机制，并为开发更高效的聚酯类降解酶提供关键指导。

2.3 作用于PHA地膜的降解微生物及酶

PHA是一类由微生物发酵合成的具有不同结构的脂肪族共聚酯，PHB、3-羟基丁酸酯和3-羟基戊酸酯的共聚物[poly(3-hydroxybutyrate-3-hydroxyvalerate), PHBV]等均属于PHA家族成员。因其良好的生物相容性和降解性，PHA可被多种微生物及其分泌酶高效降解^[31,47]。Pérez-Arauz等^[48]通过富集传代实验成功筛选出一批混合菌群，这些菌群在80 d内对PHA膜片的降解率达到75%。在这些混合菌中，假单胞菌(Pseudomonas sp.)、大肠埃希氏菌(E. coli)、盐单胞菌(Halomonas spp.)、沙克尔顿号马加利特氏菌(Margalitia shackletonii)和杀虫贪铜菌(Cupriavidus necator)等发挥了关键作用。Chhetri等^[49]从土壤中分离到一株能降解PHB的马赛菌(Massilia sp.) JJY02，该菌在28 ℃条件下仅需20 d就能对PHB实现90%的高效降解，其降解效率远高于Pérez-Arauz等^[48]筛选到的混合降解菌群。

Abe等^[50]进一步证实了微生物可通过分泌胞外解聚酶来降解PHA，并且这些微生物可利用分解的化合物作为营养物质进行生长。目前，已从海勒氏假单胞菌^[51]、施氏假单胞菌^[52]、粪产碱菌^[53]、睾丸酮丛毛单胞菌^[54]和嗜酸乳杆菌^[55]等细菌中分离纯化出多种PHB解聚酶。这些解聚酶的共性研究表明，在行使其活性功能时，至少需要识别聚合物链中的3个单体单元作为酯键水解的底物位点；其水解过程始于链表面的非结晶性聚酯部分，并逐渐深入到结晶态的聚合物链中。值得注意的是，聚酯链的序列结构对水解速率有显著影响。

PHA材料的生物降解性能与其结构特征密切相关，同时，微生物合成PHA的过程以及PHA在环境中的降解过程都与环境微生物群落组成呈现显著关联性^[56]。上述所涉及的解聚酶对聚酯链序列结构的识别机制，揭示了聚酯降解过程中单体单元排布对酶促水解的动力学影响，这为未来开发可控降解速率的BDMs提供了理论依据。

2.4 作用于PBAT地膜的降解微生物及酶

PBAT是一种由己二酸、1,4-丁二醇和对苯二甲酸缩聚合成的脂肪族-芳香族共聚酯。研究表明，PBAT地膜能被多种微生物类群降解，如芽孢杆菌属、假单胞菌属、布鲁氏菌属、肠杆菌属和微杆菌属等^[57]。特别地，一些能在地膜上定殖并富集的微生物，如节杆菌属、鞘氨醇单胞菌属、恶臭假单胞菌等，通常具备降解PBAT地膜的潜力^[58]。然而，不同微生物对PBAT的降解能力存在显著差异。刘佳茜等^[59]从环境中筛选出2株假单胞菌，Pseudomonas sp. RD1-3和Pseudomonas sp.N1-2，它们在28 ℃条件下56 d内对PBAT的降解率仅有6.88%和6.49%。Sun等^[60]分离出一株能以PBAT为唯一碳源生长的霍氏肠杆菌(Enterobacter hormaechei) WX-2，该菌在60 d内对PBAT的降解率为20%-25%。Jia等^[61]报道褐色喜热裂孢菌(Thermobifida fusca) FXJ-1在9 d内对PBAT以及PBAT-PLA复合膜的降解率可分别达到80%和50%，在证实T. fusca对PBAT具有较高降解能力的同时，却与通常认为共混物生物降解率随PBAT含量的增加而降低的理论相悖^[62]。理论上，PLA与PBAT进行复合时应当产生协同降解效应^[63]，因为PLA分子链中的易水解酯键与PBAT结构中的微生物酶解酯基可形成互补作用位点，且PLA的引入能破坏PBAT的半结晶结构有序性，改善复合体系的表面湿润性，从而促进水分子渗透和微生物定殖，进而加速降解。然而，实验数据显示，复合体系的降解速率反而低于纯PBAT体系，这可能与PLA改性导致的微区结晶重构或微生物群落的环境适应性差异有关。

PBAT在土壤中的降解还受多种环境因素制约，包括土壤条件、类型和气候条件等。Zhang等^[64]研究发现，PLA/PBAT在湿度60%和pH 7.0的条件下降解率最高，这可能与土壤细菌群落的积极响应有关；高通量测序显示，适宜的降解条件显著提升了如假单胞菌和鞘氨醇单胞菌等降解相关物种的丰度。Han等^[65]则通过土壤微宇宙系统探究PBAT地膜降解的关键因素，发现塿土中PBAT地膜的矿化率(16.0%)远高于潮土(9.0%)、黑土(0.3%)和红壤(0.9%)；宏基因组分析揭示，塿土中微生物群落对PBAT响应更敏感，且富含PBAT水解酶基因。此外，慢生根瘤菌属(Bradyrhizobium)、贪噬菌属(Variovorax)和沙壤土杆菌属(Ramlibacter)等被鉴定为新型潜在降解菌，其富集程度与PBAT降解能力呈正相关^[25,65]。这些发现强调了PBAT降解与土壤环境及细菌群落的紧密联系，凸显了这些因素在决定降解效率中的重要性。

目前已从细菌、真菌等多种微生物中获得PBAT降解酶。Dresler等^[66]从T. fusca中分离到对脂肪族-芳香族共聚酯具有超强降解能力的水解酶TfH，并发现编码该酶的基因(bta1和bta2)位于一个操纵子上，氨基酸同源性为92%。Gouda等^[67]进一步发现TfH酶在培养基中的分泌量与添加至培养基中的聚酯数量相关，属于诱导酶。Wallace等^[68]通过蛋白质组学筛选方法从食油假单胞菌(Pseudomonas oleovorans)中发掘了新的解聚酶PpEst，发现其可水解PBAT生成对苯二甲酸和4-羟基丁氧羰基苯甲酸。Yang等^[69]证实角质酶是极具潜在能力的PBAT降解酶，能够在48 h内完全分解PBAT。同时，通过应用双突变策略使底物结合口袋更具灵活性而设计的工程化角质酶，表现出更高的降解速率。这些发现增进了酶介导PBAT降解的分子理解，有助于未来的酶工程发展，以推动PBAT生物降解技术。

2.5 作用于PPC地膜的降解微生物及酶

PPC是由CO₂和环氧丙烷在催化剂作用下，通过共聚反应制备而成的高分子聚合物。PPC因其强疏水性阻碍了微生物的附着与富集，进而限制其自身的降解速率。陶剑等^[70]采用土悬法降解PPC，在第30天仍未观察到明显的降解现象。Du等^[71]进行土埋PPC地膜降解试验也证实了这一点，在第30天和第120天分别测得降解率约为1.69%和3.00%；通过摇瓶实验对PPC地膜降解，发现在pH 6.0的条件下经120 d的降解，PPC地膜的降解率达6.32%。在此期间，膜表面的亲水性显著增强，提示液体环境可能改变了膜表面的亲疏水性，从而提升了PPC地膜的降解效果。

Liang等^[72]研究了PPC覆膜对土壤理化性质和微生物群落的影响，发现与无膜条件相比，覆膜并未对土壤微生物群落及多样性产生显著影响，但促进了嗜酸杆菌的富集。张加凡^[73]通过Illumina Nova技术对覆膜各阶段的微生物群落结构进行分析，发现根瘤菌属、假单胞菌属和黄杆菌属等菌群在PPC地膜降解过程中逐渐富集。同时，该团队筛选获得了一株能以PPC为唯一碳源生长的枯草芽孢杆菌J16，在降解30 d后，PPC地膜的质量损失率达到了6.00%-9.95%^[74]。目前，PPC地膜降解菌株及其关键酶的研究仍处于起步阶段，相关报道极为稀缺。未来亟待进一步挖掘降解微生物资源，解析其代谢途径与降解酶的作用机制。

综上所述，不同类型的生物降解地膜均有其特定的有效降解微生物类群。值得关注的是，在PPC、PBAT、聚丁二酸-己二酸丁二酯(adipic acid-1,4-butanediol-succinic acid copolymer, PBSA)地膜覆盖的土壤中均富集到了含有根瘤菌属的潜在降解菌群^[25,65,75]。众所周知，根瘤菌能与豆科植物共生形成根瘤，固定并利用空气中的氮气供植物所需；现今的研究揭示，根瘤菌等固氮菌属在降解地膜方面也发挥着潜在作用，这预示着在漫长的进化历程中，根瘤菌可能演化出了复杂多样的酶系统或代谢路径^[25,65]。这些机制不仅让它们能够适应与豆科植物的共生固氮关系，还赋予了它们处理新型人工聚合物的能力。表2对作用于BDMs的部分降解微生物类群及基本特性进行了总结。

3 微生物降解BDMs的主要机制

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BDMs的降解是一个复杂而有序的生物化学过程，主要涉及微生物及其分泌酶的作用。无论真菌或细菌，降解BDMs的本质是微生物分泌的解聚合酶使其化学键断裂^[84-85]，具体途径取决于所涉及的微生物种类和聚合物类型^[24]。尽管现阶段关于BDMs的具体降解机制尚未完全阐明，但依据现有研究成果，其降解过程主要分为以下4个步骤^[86] (图1)：生物吸附附着、解聚、同化和矿化。

(1) 吸附附着阶段，微生物通过自身结构，如鞭毛、菌毛及细胞壁表面电荷等，与待降解底物发生接触并附着^[87]。这一过程的效率受多种环境因素的影响，包括温度、pH、底物浓度以及微生物的趋化性等^[84]。这些因素共同作用，引导着微生物向BDMs底物移动并有效附着。其中，细菌α多样性主要响应材料类型与温度变化，而真菌群落结构对温度梯度呈特异性响应。β多样性分析显示所有实验变量均具有显著影响，其中，Ju等^[88]通过宏基因组学证实塑料组分和时间演替对地膜表面细菌群落的β多样性具有交互作用。这些核心微生物组解析鉴定出具有潜在降解功能的优势菌属和真菌分类单元。(2) 进入解聚作用阶段，微生物分泌具有高度特异性的解聚酶，如水解酶、氧化还原酶等^[89]。基于16S rRNA基因功能，预测到水解酶、角质酶、纤维素酶和脂肪酶等关键功能基因显著富集。这些酶通过分子识别机制精准识别并结合底物化学键，通过水解、氧化等化学反应，将复杂的聚合物分解为低聚物或单体小分子。这一过程是BDMs降解的关键步骤，为后续的吸收同化奠定了基础。(3) 在吸收同化阶段，解聚产生的小分子通过跨膜运输系统(主动运输/被动扩散)进入微生物胞内代谢网络^[90]。在细胞内，这些小分子参与各类代谢途径，既可以作为碳源、氮源参与氨基酸/核苷酸等生物大分子合成，也可以经氧化磷酸化途径生成ATP，为微生物的生长、繁殖以及维持生命活动提供了必要的支持。(4) 矿化作用阶段，在微生物利用BDMs底物的过程中，部分有机物会被彻底氧化^[91]。这一过程通过三羧酸循环等途径，使有机物最终转化为CO₂和H₂O，参与到生态系统的循环之中。

值得注意的是，BDMs的降解过程还受到多种环境因素的影响。其中，温度是一个关键因素^[92]。研究表明，微生物在土壤中对塑料的有效生物降解一般发生在中温温度范围内，即10-50 ℃，这有利于微生物保持最佳活性并促进降解过程的进行。此外，塑料成分和时间演替也显著影响了地膜表面微生物群落的多样性，进而影响了BDMs的降解效率和模式。

4 BDMs对土壤生态的影响

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BDMs覆盖土壤过程中，其物理属性与普通PE地膜颇为相似，二者均能通过调节土壤微气候间接影响土壤生态系统和微生物群落功能^[93]。地膜覆盖作为一种物理屏障，显著减少了土壤的水分蒸发，抑制了气体交换，提高了土壤温度，并降低了光透过率。这些变化共同改变了土壤的理化性质^[94]。同时，土壤湿度、温度及蒸汽扩散率等参数的调整还可能进一步调节土壤微生物的活性，影响微生物的群落结构^[95] (图2)。

尽管BDMs与普通PE地膜在作为物理屏障方面具有相似性，但它们在土壤中的最终归宿却是截然不同的。生长季结束后，普通PE地膜会从土壤表面去除或以残膜形式存在，而BDMs则是在土壤中自然降解。因此，BDMs对土壤生态的影响应该与普通PE地膜存在差异。王卫星等^[96]采用覆盖对照试验，在大豆种植中验证了PBAT地膜的实际效果，发现其除草效果更佳，且能显著提升大豆的产量。赵军等^[97]对比了PBAT地膜、PE地膜覆盖和无覆盖膜3种情况下棉花的产量，结果显示，相较于PE地膜组，PBAT地膜组的棉花产量提高了13%，土壤含水量增加4.2%，且土壤盐分动态表现和水分利用效率更优，证实了BDMs通过调控水盐运移，在维持土壤健康的同时实现产量增益。在确认BDMs农艺优势的同时，其是否会出现残膜污染问题也应引起关注。Wang等^[98]研究表明，PPC、PBAT和聚二氧化碳(poly-carbon dioxide, PCO₂)这3种不同类型的BDMs在作物生长期内几乎完全降解，残留量显著低于普通PE地膜。这与齐英等^[99]的研究结果一致，即BDMs在花生收获期的残留量相较于普通地膜降低57%-67%。

BDMs是逐级分解的，在此过程中可能会形成各种降解产物。Palsikowski等^[100]以洋葱(Allium cepa)为试验对象，研究了PBAT和PLA地膜在土壤中不同自然降解阶段水提取物的生态毒理效应，结果表明各阶段水提物对A. cepa无细胞毒性、基因毒性或诱变效应。Muroi等^[101]研究了PBAT地膜自然降解过程对土壤微生物群落的影响，证实PBAT在土壤中的降解促进了真菌和根际促生菌的生长，并且对芸薹属植物无生态毒理影响，表明PBAT适宜于农业应用。Satti等^[102]利用PLA降解菌株和土著微生物对土壤进行生物增强和生物刺激，结果表明与自然降解相比，生物刺激和生物强化可显著提高PLA的矿化速率，但不影响土壤固有微生物种群。然而，也有研究表明BDMs对土壤生态的影响应予以重视。Balestri等^[103]评估了PBAT/淀粉和高密度聚乙烯(high density polyethylene, HDPE)混合物渗滤液的植物毒性效应，结果表明渗滤液对种子萌发无影响，但有相当数量的幼苗表现出发育异常或幼苗生长减少。虽然目前尚无直接证据证明BDMs在自然环境中会形成微塑料，但已有研究关注人工制备BDMs微塑料对土壤生态的潜在影响。Schöpfer等^[104]研究了PLA/PBAT和低密度聚乙烯(low density polyethylene, LDPE)植被的微塑料对秀丽隐杆线虫的潜在毒性，发现微塑料污染土壤对线虫的繁殖(土壤食物网的主要角色)有影响，并可能对土壤功能产生负面影响。Sun等^[105]研究了传统塑料PE、聚苯乙烯(polystyrene, PS)与可生物降解塑料PLA、聚丁二酸丁二醇酯(polybutylene succinate, PBS)微塑料对土壤微生物群落组成及功能的潜在影响，结果表明可生物降解塑料显著改变了土壤微生物群落结构，进而影响土壤功能。

尽管目前已从BDMs对土壤理化性质、微生物群落、作物产量、生态毒性等方面开展了相关研究，但大多局限于短期范畴，难以全面揭示BDMs的长期潜在影响。后续研究应在此基础上进行多维度、长时间的动态监测与深度分析，精准解析其在土壤生态系统中的作用机制，以突破短期研究的局限并填补该领域在长期研究方面的空白，为BDMs的环境生态风险评估和安全应用推广提供更为科学和全面的依据。

5 总结与展望

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生物可降解地膜作为传统地膜最具潜力的替代品，正展现出巨大的市场潜力和环保价值。为促进具有我国自主知识产权的生物可降解地膜得以广泛应用，深入探究具有共性结构的聚酯和聚碳酸酯类生物可降解地膜的降解机制及生态环境效应至关重要。本文聚焦于5种极具应用潜力的生物可降解地膜，系统概述了它们的基本特性，并深入分析了其降解微生物类群及关键降解酶在降解过程中的作用机制。然而，当前研究更关注于生物可降解地膜对农作物产量的提升，对其降解机制及生态环境效应的探讨尚显不足。鉴于生物可降解地膜推广规模的不断扩大，未来研究需着重于以下关键内容。

(1) 高效地膜降解微生物的挖掘与关键酶的解析。通过先进技术手段，精准解析酶与底物的相互作用模式，优化现有降解酶的设计，实现定向改造，从而明确生物可降解地膜降解的限速步骤，提升降解的可控性。

(2) 明确生物可降解地膜逐级降解过程的中间产物及其生态影响。深入分析逐级降解中间产物的积累特征，阐明逐级分解途径，评估其对生态环境的短期与长期潜在影响。

(3) 关注生物可降解地膜微塑料形成及其环境效应。明确生物可降解地膜微塑料形成的可能性，评估人工制备的各类生物可降解地膜微塑料的潜在生态影响。

这些研究将为生物可降解地膜的大规模安全应用、降解的可调控性的实现奠定基础。通过科学评估与合理调控，确保生物可降解地膜在促进农业生产的同时，不会对生态环境造成不可逆的损害，最终实现农业可持续发展和生态环境保护的双赢目标。

致谢

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感谢在线平面设计网站(gdp.renlab.cn)提供绘图元素。

作者利益冲突公开声明

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作者声明不存在任何可能会影响本文所报告工作的已知经济利益或个人关系。

基金

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国家重点研发计划(2018YFD1001000)
陕西省秦岭生态安全重点实验室开放基金(QLES202501)
中国科学院西安分院“西部之光”人才培养计划(2022S-1)
山西省重点研发计划(202102140601011)

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2025年第65卷第6期

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doi: 10.13343/j.cnki.wsxb.20250044

接收时间：2025-01-15
首发时间：2026-02-07
出版时间：2025-06-04

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收稿日期：2025-01-15
录用日期：2025-03-16

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National Key Research and Development Program of China(2018YFD1001000)

国家重点研发计划(2018YFD1001000)

Key Laboratory Project of Shaanxi Provincial of Qinling Ecological Security(QLES202501)

陕西省秦岭生态安全重点实验室开放基金(QLES202501)

“Light of the West” Talent Cultivation Program of Xi’an Branch of the Chinese Academy of Sciences(2022S-1)

中国科学院西安分院“西部之光”人才培养计划(2022S-1)

Key Research and Development Program of Shanxi Province(202102140601011)

山西省重点研发计划(202102140601011)

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^1.西安建筑科技大学环境与市政工程学院，陕西西安

^2.陕西省微生物研究所，陕西西安

^3.陕西省秦岭生态安全重点实验室，陕西西安

^4.山西农业大学农业经济管理学院，山西太原

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2种不同金属材料的力学参数

科 Family	属数 Number of genus	种数 Number of species	占总种数比例 Percentage of total species (%)	属 Genus	种数 Number of species	占总种数比例 Percentage of total species (%)
鹅膏菌科Amanitaceae	2	11	5.26	鹅膏菌属 Amanita	10	4.78
小菇科 Mycenaceae	2	12	5.74	丝盖伞属 Inocybe	5	2.39
多孔菌科 Polyporaceae	8	14	6.70	蜡蘑属 Laccaria	5	2.39
红菇科 Russulaceae	3	23	11.00	小皮伞属 Marasmius	6	2.87
				小菇属 Mycena	11	5.26
				光柄菇属 Pluteus	5	2.39
				红菇属 Russula	17	8.13
				栓菌属 Trametes	5	2.39

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TxT

材料 Material	单体 Monomer	常见聚合方式 Common polymerization methods	物理性能 Physical performance	参考文献References
Polycaprolactone (PCL)	己内酯 ε-caprolactone	Open-loop polymerization method	Exhibits depressed thermal transitions with a melting point (T_m=60 ℃) and glass transition temperature (T_g=-60 ℃), demonstrating exceptional processability	[14]
Polylactic acid (PLA)	乳酸(或丙交酯) Lactic acid	Polymerization, open-loop polymerization method, enzymatic polymerization	Distinctive thermal stability manifested through elevated T_m (170 ℃) and T_g (60 ℃) exceeding conventional aliphatic polyester systems	[15]
Polyhydroxyalkanoates(PHA)	多种羟基脂肪酸 Hydroxyalkanoic acid	-	Presents a T_mof 145 ℃ with adjustable physicochemical properties through monomer compositional engineering and stoichiometric optimization	[16]
Poly(β- hydroxybutyrate) (PHB)	3-羟基丁酸 3-hydroxybutyric acid	Displays substantially elevated T_m(180 ℃) concomitant with superior mechanical rigidity and pronounced brittle fracture behavior	[17-18]
Poly(3-hydroxybutyrate-3-hydroxyvalerate (PHBV)	3-羟基丁酸 3-hydroxybutyric acid 3-羟基戊酸 3-hydroxyvaleric acid	Enables precise property tailoring via controlled 3-hydroxyvalerate content modulation in copolymer architecture	[19]
Poly(butylene adipate-co-terephthalate (PBAT)	己二酸 Adipic acid 对苯二甲酸Terephthalic acid 1,4-丁二醇 1,4-butanediol	Melt polycondensation	Thermal profile comprises: T_m= 110-130 ℃, T_g=-30 ℃, crystallization temperature (T_c) 110 ℃, and crystallinity degree (X_c) 30%	[20-21]
Poly propylene carbonate (PPC)	二氧化碳Carbon dioxide 环氧丙烷Propylene oxide	Coordination copolymerization	Features markedly elevated phase-transition temperatures with T_m=160-170 ℃ and T_g=30-41 ℃	[22]

材料

Material

单体

Monomer

结构式

Structural formulas

常见聚合方式

Common polymerization methods

物理性能

Physical performance

参考文献References

Polycaprolactone (PCL)

己内酯

ε-caprolactone

Open-loop polymerization method

Exhibits depressed thermal transitions with a melting point (T_m=60 ℃) and glass transition temperature (T_g=-60 ℃), demonstrating exceptional processability

[14]

Polylactic acid (PLA)

乳酸(或丙交酯)

Lactic acid

Polymerization, open-loop polymerization method, enzymatic polymerization

Distinctive thermal stability manifested through elevated T_m (170 ℃) and T_g (60 ℃) exceeding conventional aliphatic polyester systems

[15]

Polyhydroxyalkanoates(PHA)

多种羟基脂肪酸

Hydroxyalkanoic acid

Presents a T_mof 145 ℃ with adjustable physicochemical properties through monomer compositional engineering and stoichiometric optimization

[16]

Poly(β- hydroxybutyrate) (PHB)

3-羟基丁酸

3-hydroxybutyric acid

Displays substantially elevated T_m(180 ℃) concomitant with superior mechanical rigidity and pronounced brittle fracture behavior

[17-18]

Poly(3-hydroxybutyrate-3-hydroxyvalerate (PHBV)

3-羟基丁酸

3-hydroxybutyric acid

3-羟基戊酸

3-hydroxyvaleric acid

Enables precise property tailoring via controlled 3-hydroxyvalerate content modulation in copolymer architecture

[19]

Poly(butylene adipate-co-terephthalate (PBAT)

己二酸

Adipic acid

对苯二甲酸Terephthalic acid

1,4-丁二醇

1,4-butanediol

Melt polycondensation

Thermal profile comprises: T_m=

110-130 ℃, T_g=-30 ℃, crystallization temperature (T_c) 110 ℃, and crystallinity degree (X_c) 30%

[20-21]

Poly propylene carbonate (PPC)

二氧化碳Carbon dioxide

环氧丙烷Propylene oxide

Coordination copolymerization

Features markedly elevated phase-transition temperatures with T_m=160-170 ℃ and T_g=30-41 ℃

[22]

材料 Material	生物可降解地膜类型 Types of biodegradable mulch film	菌株 Strain	菌株来源 Source	降解周期 Degradation period (d)	降解率 Degradation rate	参考文献References
PCL	PCL mulch film	Pseudomonas sp. DS0901	Activated sludge	3	100%	[32]
PLA	PLA mulch film	Pseudomonas geniculata WS3	Soil and wastewater sludge	20	Around 45%	[42]
PLA mulch film	Stenotrophomonas pavanii CH1	/	20	10%	[42]
PLA mulch film	Bacillus pumilus B12	Soil	2	/	[76]
PLA powder/tray	Actinomadura keratinilytica T16-1	/	3	99%/32%	[77]
PLA mulch film	Nocardiopsis prasina	Ocean	60	(1.27±0.19)%	[78]
PLA mulch film	Priestia flexa PMPHB7	Soil	21	73%	[79]
PLA mulch film	Bacillus safensis PLA1006	Soil	30	8%	[80]
PLA mulch film	Tritirachium album	/	4	78%	[81]
PHA	PHA mulch film	/	Soil	80	75%	[48]
PHB	PHB mulch film	Acinetobacter junii BP25	Wastewater	77	83%	[82]
PHBV	PHBV	Alcaligenes spp. Pseudomonas spp.	Soil	365	13%	[83]
PBAT	PBAT mulch film	Enterobacter hormaechei WX-2	Soil	60	(20.8±2.2)%	[60]
PBAT mulch film (PF, AF, PPSF)	Thermobifida fusca FXJ-1	Compost samples	9	(82.87±1.01)%, (87.83±2.00)%, (52.53±0.54)%	[61]
PPC	PPC mulch film	Bacillus subtilis J16	Soil	30	9.95%	[73]

材料

Material

生物可降解地膜类型

Types of biodegradable mulch film

菌株

Strain

菌株来源

Source

降解周期

Degradation period (d)

降解率

Degradation rate