Bacterial Community Characteristics during the Mixed Decomposition of Litter from Pinus massoniana and Indigenous Broad-leaved Tree Species in Southwestern China

doi:10.16258/j.cnki.1674-5906.2024.01.002

Ecology and Environment ›› 2024, Vol. 33 ›› Issue (1): 12-27.DOI: 10.16258/j.cnki.1674-5906.2024.01.002

• Research Article • Previous Articles Next Articles

Bacterial Community Characteristics during the Mixed Decomposition of Litter from Pinus massoniana and Indigenous Broad-leaved Tree Species in Southwestern China

LI Xun¹(), ZHANG Yan¹, SONG Simeng¹, ZHOU Yang¹, ZHANG Jian²^,^*()

1. Research Center for Ecological Restoration and Characteristic Industry Cultivation in Hengduan Mountains Region/Research Center of Forest-Grass Care and Use in Sichuan-Tibet-Yunnan-Qinghai, Sichuan Minzu College, Kangding 626001, P. R. China
2. Key Laboratory of Forestry Ecological Engineering in Sichuan/Collaborative Innovation Center of Ecological Security in the Upper Reaches of Yangtze River, Sichuan Agricultural University, Chengdu 611130, P. R. China

Received:2023-04-21 Online:2024-01-18 Published:2024-03-19
Contact: ZHANG Jian

西南地区马尾松与乡土阔叶树种凋落叶混合分解过程中的细菌群落特征

李勋¹(), 张艳¹, 宋思梦¹, 周扬¹, 张健²^,^*()

1.四川民族学院横断山区生态修复与特色产业培育研究中心/川藏滇青林草抚育和利用研究中心，四川康定 626001
2.四川农业大学林学院生态林业工程重点实验室/长江上游生态安全协同创新中心，四川成都 611130

通讯作者: 张健
作者简介:李勋（1990年生），男，副教授，博士，主要从事长江上游马尾松低效林改造研究工作。E-mail: 502780405@qq.com
基金资助:
四川省科技计划项目(2023NSFSC1168);国家自然科学基金项目(31370628);甘孜州科技计划项目(220030)

Abstract

Abstract:

Tree species diversity in forest ecosystems not only controls the diversity and proportion of litter but also affects the microbial community residing in decomposed litter. However, the differences in the bacterial community structure in the process of leaf litter decomposition with different tree species combinations and mixed proportions are still unclear. In this study, a mixture of litter from Pinus massoniana and native broad-leaved tree species, including Toona sinensis (A. Juss.) Roem., Cinnamomum camphora Linn., and Sassafras tzumu Hemsl., were created based on various tree species and their corresponding mass proportions. This study utilized Illumina MiSeq high-throughput sequencing to examine the dynamic features of the bacterial community structure during leaf litter decomposition. Proteobacteria and Actinobacteria were the dominant bacterial groups in all the treatments. The dominant genera were Sphingomonas, unidentified Rhizobiaceae, Bradyrhizobium, and unidentified Cyanobacteria. Overall, the diversity and richness of the bacteria in the mixed litter showed a strong synergistic effect after 250 days of decomposition (70.97% and 29.03%, respectively), whereas during the second year of decomposition, most of the observed values for the mixed litter were less than 0. Especially in the late decomposition stage, the bacterial diversity index of 19.35% mixed litter and bacterial richness index of 9.68% mixed litter showed antagonistic effects. Among different tree species combinations, the combination of P. massoniana +C. camphora +T. sinensis (PCT) with a broad-leaved ratio of 30%‒40% is more conducive to improving the diversity and relative richness of bacterial species in leaf litter. In summary, the diversity and richness index of bacteria in leaf litter were comprehensively influenced by the decomposition period and tree species combination. In the later stage of the transformation of low-efficiency artificial pure P. massonianaforests, C. camphora and T. sinensis could be planted to transform monoculture P. massoniana forests into mixed forests.

Key words: mixed litter, bacterial, Pinus massoniana, high-throughput sequencing, non-additive effect

摘要：

林地内凋落叶的种类和比例调控其分解过程中微生物的群落结构。然而，不同树种组合以及不同混合比例的凋落叶在分解过程中细菌群落结构有何差异，目前尚不清楚。将马尾松与乡土阔叶树种香椿[Toona sinensis (A. Juss.) Roem.]、香樟（Cinnamomum camphora Linn.）和檫木（Sassafras tzumu Hemsl.）凋落叶按照不同树种、不同质量比例混合后，通过Illumina MiSeq高通量测序研究凋落叶分解过程中细菌群落结构的动态变化特征。结果表明，在所有处理中，Proteobacteria和Actinobacteria均为优势门，Sphingomonas，unidentified_Rhizobiaceae，Bradyrhizobium和unidentified_Cyanobacteria为优势菌属。此外，混合凋落叶中细菌的多样性和丰富度在分解250 d后表现出较强的协同效应（分别为70.97%和29.03%）；第2年分解期内大部分混合凋落叶的观测值-期望值<0，尤其是分解末期（分解604 d）后有19.35%的混合凋落叶的细菌多样性指数和9.68%的混合凋落叶的细菌丰富度指数表现出拮抗效应。不同树种组合凋落叶中，马尾松+香樟+香椿组合（PCT）下阔叶占比为30%-40%时更有利于提高细菌的多样性和丰富度。综上，凋落叶中细菌的多样性和丰富度受到分解时期和树种组合的共同影响。在后期马尾松纯林的“混交化”改造过程中，适当引入乡土阔叶树种香樟与香椿可增加马尾松林地内凋落叶中细菌的多样性和类群丰度。

关键词: 混合凋落叶, 细菌, 马尾松, 高通量测序, 非加性效应

CLC Number:

LI Xun, ZHANG Yan, SONG Simeng, ZHOU Yang, ZHANG Jian. Bacterial Community Characteristics during the Mixed Decomposition of Litter from Pinus massoniana and Indigenous Broad-leaved Tree Species in Southwestern China[J]. Ecology and Environment, 2024, 33(1): 12-27.

李勋, 张艳, 宋思梦, 周扬, 张健. 西南地区马尾松与乡土阔叶树种凋落叶混合分解过程中的细菌群落特征[J]. 生态环境学报, 2024, 33(1): 12-27.

Figures/Tables 15

References 54

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科	属	种	生活型
豆科 Fabaceae	胡枝子属 Lespedeza	胡枝子 Lespedeza bicolor	灌木
芸香科 Rutaceae	花椒属 Zanthoxylum	野花椒 Zanthoxylum simulans	灌木
蔷薇科 Rosaceae	悬钩子属 Rubus	针刺悬钩子 Rubus pungens	灌木
海桐科 Pittosporaceae	海桐属 Pittosporum	光叶海桐 Pittosporum glabratum	灌木
小檗科 Berberidaceae	十大功劳属 Mahonia	十大功劳 Mahonia fortunei	灌木
莎草科 Cyperaceae	莎草属 Cyperus	莎草 Cyperus rotundus	草本
碗蕨科 Dennstaedtiaceae	蕨属 Pteridium	欧洲蕨 Pteridium aguilinum	草本
天门冬科 Asparagaceae	沿阶草属 Ophiopogon	麦冬 Ophiopogon japonicus	草本
羽藓科 Thuidiaceae	山羽藓属 Abietinella	山羽藓 Thuidium assimile	草本
鸢尾科 Iridaceae	鸢尾属 Iris	蝴蝶花 Iris japonica	草本
里白科 Gleicheniaceae	芒萁属 Dicranopteris	芒萁 Dicranopteris pedata	草本

科	属	种	生活型
豆科 Fabaceae	胡枝子属 Lespedeza	胡枝子 Lespedeza bicolor	灌木
芸香科 Rutaceae	花椒属 Zanthoxylum	野花椒 Zanthoxylum simulans	灌木
蔷薇科 Rosaceae	悬钩子属 Rubus	针刺悬钩子 Rubus pungens	灌木
海桐科 Pittosporaceae	海桐属 Pittosporum	光叶海桐 Pittosporum glabratum	灌木
小檗科 Berberidaceae	十大功劳属 Mahonia	十大功劳 Mahonia fortunei	灌木
莎草科 Cyperaceae	莎草属 Cyperus	莎草 Cyperus rotundus	草本
碗蕨科 Dennstaedtiaceae	蕨属 Pteridium	欧洲蕨 Pteridium aguilinum	草本
天门冬科 Asparagaceae	沿阶草属 Ophiopogon	麦冬 Ophiopogon japonicus	草本
羽藓科 Thuidiaceae	山羽藓属 Abietinella	山羽藓 Thuidium assimile	草本
鸢尾科 Iridaceae	鸢尾属 Iris	蝴蝶花 Iris japonica	草本
里白科 Gleicheniaceae	芒萁属 Dicranopteris	芒萁 Dicranopteris pedata	草本

样地	pH	w_(全碳)/ (g∙kg⁻¹)	w_(全氮)/ (g∙kg⁻¹)	土壤容重/ (g∙cm⁻³)	海拔高度/ m	坡度/ (°)	坡向
1	4.60± 0.20	12.3± 2.01	0.73± 0.15	1.42± 0.03	811± 13.4	10.7± 5.31	S
2	4.10± 0.10	13.7± 2.11	0.70± 0.11	1.41± 0.11	825± 11.5	13.1± 6.31	SE
3	4.10± 0.10	14.1± 3.12	0.70± 0.21	1.42± 0.14	812± 12.4	16.2± 4.06	SE

样地	pH	w_(全碳)/ (g∙kg⁻¹)	w_(全氮)/ (g∙kg⁻¹)	土壤容重/ (g∙cm⁻³)	海拔高度/ m	坡度/ (°)	坡向
1	4.60± 0.20	12.3± 2.01	0.73± 0.15	1.42± 0.03	811± 13.4	10.7± 5.31	S
2	4.10± 0.10	13.7± 2.11	0.70± 0.11	1.41± 0.11	825± 11.5	13.1± 6.31	SE
3	4.10± 0.10	14.1± 3.12	0.70± 0.21	1.42± 0.14	812± 12.4	16.2± 4.06	SE

树种丰富度	树种组合	缩写	混合比例
1	马尾松(P. massoniana)、檫木(S. tsumu)、香樟(C. camphor)、香椿(T. sinensis)	P、S、C、T	10꞉0
2	马尾松+香樟 P. massoniana+C. camphor	PC	8꞉2、7꞉3、6꞉4
	马尾松+檫木 P. massoniana+S. tsumu	PS
	马尾松+香椿 P. massoniana+T. sinensis	PT
3	马尾松+檫木+香樟 P. massoniana+S. tsumu+C. camphor	PSC	8꞉1꞉1、7꞉2꞉1、7꞉1꞉2、6꞉2꞉2、6꞉3꞉1、6꞉1꞉3
	马尾松+香樟+香椿 P. massoniana+C. camphor+T. sinensis	PCT
	马尾松+檫木+香椿 P. massoniana+S. tsumu+T. sinensis	PST
4	马尾松+檫木+香樟+香椿 P. massoniana+S. tsumu+C. camphor+T. sinensis	PSCT	7꞉1꞉1꞉1、6꞉2꞉1꞉1、6꞉1꞉1꞉2、6꞉1꞉2꞉1

Bacterial Community Characteristics during the Mixed Decomposition of Litter from Pinus massoniana and Indigenous Broad-leaved Tree Species in Southwestern China

西南地区马尾松与乡土阔叶树种凋落叶混合分解过程中的细菌群落特征

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混合处理	C	N	P	木质素	纤维素	总酚	单宁
PT82	437.96±8.90ab	7.15±0.42abc	1.02±0.03abcdefghi	307.74±11.27abc	129.01±18.71a	49.15±2.77abc	20.54±1.06bcd
PT73	430.58±7.95ab	7.68±0.28abc	1.07±0.03abc	286.07±9.61abcde	125.28±17.06a	46.63±2.34abcde	18.43±0.93def
PT64	423.20±7.05b	8.22±0.14a	1.12±0.04a	264.40±8.19e	121.55±15.42a	44.12±1.91abcdef	16.32±0.80f
PS82	444.92±8.17a	6.13±0.65c	0.91±0.03jkl	315.46±11.76a	128.54±16.63a	50.42±2.92a	64.94±0.34a
PS73	441.02±6.86ab	6.16±0.63bc	0.91±0.03kl	297.65±10.70abcd	124.57±13.98a	48.55±2.57abc	66.04±0.35a
PS64	437.12±5.62ab	6.19±0.62bc	0.90±0.03l	279.85±10.15b	120.61±11.39a	46.67±2.21abcde	67.13±0.39a
PC82	446.32±6.50a	6.50±0.71abc	0.96±0.03efghijkl	310.78±11.88ab	138.11±16.48a	46.30±3.00abcdef	22.48±1.05b
PC73	443.13±4.32a	6.71±0.71abc	0.98±0.04cdefghijkl	290.64±10.43abcde	138.94±13.80a	42.37±2.69bcdef	21.34±0.91bc
PC64	439.93±2.14ab	6.93±0.72abc	1.00±0.05bcdefghijkl	270.5±9.08de	139.77±11.24a	38.43±2.38f	20.21±0.77bcd
PST811	441.44±8.53ab	6.64±0.54abc	0.96±0.03defghijkl	311.6±11.50a	128.77±17.67a	49.78±2.84ab	21.61±0.99bc
PST721	437.54±7.23ab	6.67±0.51abc	0.96±0.03efghijkl	293.79±10.30abcde	124.81±15.00a	47.91±2.49abcd	20.57±0.78bcd
PST712	434.06±7.59ab	7.18±0.40abc	1.01±0.03bcdefghhi	289.93±9.93abcde	125.05±16.02a	47.27±2.41abcde	19.50±0.86cde
PST631	433.64±5.97ab	6.70±0.50abc	0.96±0.03fghijkl	275.98±9.61de	120.85±12.37a	46.03±2.14abcdef	19.53±0.58cde
PST622	430.16±6.33ab	7.21±0.38abc	1.01±0.03bcdefghij	272.12±9.10de	121.08±13.38a	45.4±2.06abcdef	18.46±0.65def
PST613	426.68±6.69ab	7.71±0.26abc	1.06±0.04abcd	268.26±8.63de	121.32±14.39a	44.76±1.98abcdef	17.39±0.73ef
PSC811	445.62±7.33a	6.31±0.67bc	0.94±0.03ghijkl	313.12±11.57a	133.33±16.50a	48.36±2.96abcd	22.58±0.98b
PSC721	441.72±6.00ab	6.34±0.64bc	0.93±0.03hijkl	295.31±10.04abcde	129.36±13.78a	46.49±2.61abcde	21.54±0.77bc
PSC712	442.42±5.15ab	6.53±0.67abc	0.95±0.03fghijkl	292.98±9.95abcde	134.15±13.72a	44.43±2.65abcdef	21.44±0.84bc
PSC631	437.83±4.74ab	6.37±0.62bc	0.93±0.03ijkl	277.51±8.96cde	125.40±11.10a	44.61±2.25abcdef	20.50±0.57bcd
PSC622	438.53±3.86ab	6.56±0.64abc	0.95±0.03fghijkl	275.17±8.33de	130.19±10.97a	42.55±2.29abcdef	20.40±0.64bcd
PSC613	439.23±2.99ab	6.74±0.68abc	0.97±0.04cdefghijkl	272.83±8.37de	134.98±11.02a	40.49±2.34def	20.30±0.70bcd
PCT811	442.14±7.69ab	6.82±0.57abc	0.99±0.02bcdefghijkl	309.26±11.45ab	133.56±17.56a	47.73±2.89abcde	21.51±1.06bc
PCT721	438.95±5.52ab	7.04±0.57abc	1.01±0.02bcdefghij	289.12±9.87abcde	134.39±14.80a	43.79±2.57abcdef	20.37±0.92bcd
PCT712	434.76±6.73ab	7.36±0.43abc	1.04±0.02abcdef	287.59±9.60abcde	129.83±15.89a	45.21±2.46abcdef	19.40±0.92cde
PCT631	435.75±3.36ab	7.25±0.58abc	1.03±0.03abcdefgh	268.97±8.36de	135.21±12.12a	39.86±2.26ef	19.23±0.78cde
PCT622	431.57±4.58ab	7.58±0.43abc	1.06±0.01abcde	267.45±7.95de	130.66±13.12a	41.28±2.14cdef	18.26±0.79def
PCT613	427.39±5.82ab	7.90±0.29ab	1.09±0.02ab	265.93±7.89e	126.11±14.23a	42.70±2.03abcdef	17.29±0.79ef
PSCT7111	438.24±6.37ab	6.85±0.53abc	0.98±0.02cdefghijkl	291.45±9.79abcde	129.6±14.83a	45.85±2.53abcdef	20.47±0.85bcd
PSCT6211	434.35±5.10ab	6.88±0.51abc	0.98±0.02cdefghijkl	273.65±8.56de	125.63±12.13a	43.97±2.18abcdef	19.43±0.64cde
PSCT6121	435.05±4.22ab	7.07±0.53abc	1.00±0.02bcdefghijk	271.31±8.11de	130.42±12.04a	41.91±2.22bcdef	19.33±0.71cde
PSCT6112	430.87±5.46ab	7.39±0.39abc	1.03±0.02abcdefg	269.79±8.20de	125.87±13.17a	43.34±2.10abcdef	18.36±0.72def

分类水平	不同分解时间下细菌的16S rDNA序列
分类水平	75 d	250 d	442 d	604 d
门	48	44	43	52
纲	60	57	53	60
目	151	131	130	137
科	299	269	268	277
属	866	709	714	721

处理	OTUs	Shannon指数	Chao1指数	ACE指数
P	2501±111abcde	8.21±0.76abcd	2571±17abcde	2658±32ab
S	2638±891abcde	8.29±1.13abcd	2700±858abcde	2802±896ab
C	2675±818abcde	8.24±0.97abcd	2670±758abcde	2770±787ab
T	2714±881abcd	8.58±0.91abc	2722±806abcde	2795±827ab
PT82	2425±53de	8.38±0.3abcd	2580±111abcde	2623±117ab
PT73	2603±181abcde	8.05±0.13bcd	2662±151abcde	2724±138ab
PT64	2629±103abcde	8.38±0.16abcd	2609±155abcde	2718±153ab
PS82	2674±76abcde	8.67±0.4abc	2821±191abcd	2886±181ab
PS73	2650±109abcde	8.64±0.18abc	2671±78abcde	2751±72ab
PS64	2656±218abcde	8.75±0.17ab	2727±147abcde	2798±168ab
PC82	2551±115abcde	8.08±0.43bcd	2579±99abcde	2704±68ab
PC73	2522±118abcde	8.24±0.16abcd	2523±180abcde	2622±161abc
PC64	2369±128e	8.06±0.32bcd	2411±58de	2510±73c
PST811	2682±107abcde	8.57±0.17abc	2797±179abcde	2883±149ab
PST721	2722±327abcd	8.67±0.49abc	2801±315abcde	2884±318ab
PST712	2452±135bcde	8.25±0.23abcd	2547±76abcde	2626±68ab
PST631	2632±185abcde	8.4±0.58abcd	2725±135abcde	2834±157ab
PST622	2580±192abcde	8.07±0.38bcd	2599±169abcde	2694±146ab
PST613	2442±675cde	8.19±0.96abcd	2500±639bcde	2588±657abc
PSC811	2624±361abcde	8.23±0.87abcd	2678±349abcde	2738±389ab
PSC721	2500±144abcde	8.01±0.35cd	2401±48de	2503±21c
PSC712	2562±221abcde	8.18±0.64abcd	2705±173abcde	2774±164ab
PSC631	2745±391abc	8.27±0.61abcd	2743±320abcde	2871±351ab
PSC622	2693±117abcd	8.37±0.33abcd	2620±142abcde	2714±150ab
PSC613	2631±197abcde	8.18±0.15abcd	2539±96abcde	2632±117ab
PCT811	2516±294abcde	8.3±0.59abcd	2432±287cde	2514±302bc
PCT721	2762±197ab	8.36±0.15abcd	2872±264ab	2953±259a
PCT712	2780±150a	8.33±0.17abcd	2847±164abc	2965±196a
PCT631	2589±206abcde	8.07±0.29bcd	2763±300abcde	2871±300ab
PCT622	2752±326abc	8.84±0.54a	2865±332ab	2942±331ab
PCT613	2714±240abcd	8.69±0.39abc	2924±189a	3016±212a
PSCT7111	2698±244abcd	8.67±0.53abc	2779±228abcde	2885±272ab
PSCT6211	2423±217de	7.74±0.93d	2384±98e	2485±96c
PSCT6121	2614±169abcde	8.36±0.24abcd	2680±50abcde	2798±74ab
PSCT6112	2575±323abcde	8.51±0.41abc	2608±345abcde	2709±340ab

拉丁学名	不同分解时间细菌群落纲水平的相对丰度/%
拉丁学名	75 d	250 d	442 d	604 d
Alphaproteobacteria	43.71	42.03	33.79	32.47
Gammaproteobacteria	23.62	14.80	14.41	14.23
Bacteroidia	10.06	11.22	3.11	3.24
unidentified_Actinobacteria	9.66	12.67	17.35	18.00
Deltaproteobacteria	3.81	7.28	4.30	4.29
Acidobacteriia	2.46	1.52	2.42	2.28
unidentified_Cyanobacteria	0.83	2.08	8.46	8.46
Others	5.85	8.40	16.16	17.03

处理	变形菌门	放线菌门	鞘氨醇单胞菌属	未鉴定蓝细菌	未鉴定根瘤菌	慢生根瘤菌属
P	0.61±0.10abc	0.20±0.07abc	7.00±0.63abcd	1.90±0.71c	2.50±0.43ef	5.10±0.29abcd
S	0.64±0.10abc	0.15±0.09abc	4.80±1.46defg	1.00±0.54c	5.20±1.08b	2.00±0.25i
C	0.60±0.06abc	0.17±0.03abc	4.90±0.50defg	3.90±3.59abc	6.80±2.34a	1.60±0.30i
T	0.58±0.07abc	0.17±0.08abc	4.60±0.84efg	2.30±0.81c	4.40±0.57bcd	1.40±0.03i
PT82	0.55±0.08bc	0.23±0.06a	5.30±0.90bcdefg	6.50±3.38abc	2.70±0.16ef	5.00±0.31abcde
PT73	0.53±0.07c	0.20±0.05abc	5.40±0.83bcdefg	11.80±6.22a	3.40±0.49cde	4.70±0.47bcdefg
PT64	0.55±0.08bc	0.19±0.01abc	5.20±0.71cdefg	8.80±7.57abc	2.70±0.47ef	4.80±0.59abcdef
PS82	0.60±0.08abc	0.20±0.05abc	7.60±1.08ab	1.60±0.84c	2.30±0.10ef	6.00±0.70ab
PS73	0.55±0.11bc	0.21±0.08abc	6.10±0.26abcdef	3.90±2.36abc	2.90±0.19ef	5.80±0.35ab
PS64	0.58±0.08abc	0.20±0.06abc	7.300±1.56abc	2.10±0.82c	2.40±0.10ef	6.00±0.91ab
PC82	0.56±0.15bc	0.20±0.07abc	4.30±0.47fg	9.80±6.60abc	3.40±0.13de	5.30±1.07abc
PC73	0.56±0.15abc	0.21±0.08abc	5.40±0.39bcdefg	6.80±3.76abc	4.30±0.15bcd	5.90±0.20ab
PC64	0.54±0.15bc	0.22±0.07ab	4.60±0.54efg	9.50±9.10abc	4.60±0.94bc	4.20±0.77cdefgh
PST811	0.62±0.08abc	0.17±0.04abc	6.70±0.49abcde	4.10±3.76abc	2.90±0.27ef	5.30±0.25abc
PST721	0.62±0.07abc	0.20±0.07abc	6.60±0.43abcdef	1.40±0.47c	2.30±0.10ef	6.00±0.46ab
PST712	0.58±0.09abc	0.19±0.06abc	7.00±1.25abcde	6.50±3.54abc	2.60±0.38ef	5.60±0.27ab
PST631	0.61±0.05abc	0.19±0.08abc	5.40±0.31bcdefg	2.70±1.06bc	2.60±0.12ef	6.20±0.87a
PST622	0.58±0.13abc	0.20±0.09abc	4.90±0.47defg	8.30±1.04abc	3.20±0.18de	5.60±0.32ab
PST613	0.61±0.09abc	0.21±0.07abc	5.20±0.4cdefg	3.70±2.72abc	3.20±0.66de	4.20±0.47cdefgh
PSC811	0.65±0.06ab	0.17±0.06abc	6.10±0.56abcdef	1.70±1.18c	2.30±0.24ef	5.90±0.29ab
PSC721	0.60±0.11abc	0.21±0.10abc	7.30±0.38abc	5.90±5.70abc	3.20±0.22de	6.00±0.61ab
PSC712	0.62±0.12abc	0.19±0.09abc	7.30±1.33abc	3.50±3.08abc	2.60±0.09ef	5.40±0.44abc
PSC631	0.61±0.08abc	0.20±0.06abc	7.40±0.41abc	4.40±3.35abc	1.70±0.22f	5.60±0.59ab
PSC622	0.65±0.06ab	0.18±0.07abc	7.30±1.03abc	3.00±1.83abc	2.90±0.30ef	5.80±0.29ab
PSC613	0.60±0.10abc	0.19±0.08abc	6.60±0.15abcdef	6.70±3.48abc	2.80±0.06ef	5.60±0.65ab
PCT811	0.64±0.08abc	0.17±0.06abc	7.90±0.15a	4.60±2.88abc	3.10±0.30def	4.0±0.51defgh
PCT721	0.59±0.10abc	0.19±0.07abc	6.00±0.58abcdef	7.80±1.79abc	3.40±0.19de	3.9±0.14defgh
PCT712	0.59±0.12abc	0.18±0.08abc	6.40±1.99abcde	6.90±4.33abc	2.90±0.23ef	4.10±0.12cdefgh
PCT631	0.53±0.10c	0.19±0.03abc	5.50±0.28bcdefg	11.20±3.79bc	3.60±0.74cde	3.40±0.2gh
PCT622	0.60±0.07abc	0.17±0.05abc	5.80±0.66bcdefg	1.80±0.72c	3.20±0.09de	3.70±0.24defgh
PCT613	0.68±0.06a	0.13±0.06c	6.00±1.59abcdef	2.00±2.00c	2.70±0.14ef	3.60±0.49efgh
PSCT7111	0.65±0.08ab	0.14±0.06bc	4.30±1.0fg	1.10±0.52c	2.50±0.35ef	3.50±0.51fgh
PSCT6211	0.61±0.13abc	0.18±0.07abc	3.70±0.35g	7.30±0.29abc	2.50±0.34ef	4.80±1.58abcdefg
PSCT6121	0.61±0.06abc	0.18±0.05abc	5.30±0.79cdefg	5.60±3.47abc	2.90±0.36ef	4.20±0.17cdefgh
PSCT6112	0.63±0.04abc	0.15±0.05abc	6.50±2.41abcdef	3.30±3.69abc	3.20±0.61de	3.20±1.05h

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