Effects of Funneliformis mosseae Inoculation on the Growth and Rhizosphere Microenvironment of Sand-Cultivated Pepper Amended with Biochar

doi:10.16420/j.issn.0513-353x.2025-1284

Acta Horticulturae Sinica ›› 2026, Vol. 53 ›› Issue (4): 1073-1087.doi: 10.16420/j.issn.0513-353x.2025-1284

• Research on Arbuscular Mycorrhizal Fungi in Horticultural Plants • Previous Articles Next Articles

Effects of Funneliformis mosseae Inoculation on the Growth and Rhizosphere Microenvironment of Sand-Cultivated Pepper Amended with Biochar

YANG Yuqi, WEI Yuxin, LI Xuezhen, KE Xixi, HUANG Tingting, LIU Huiying^*()

Key Laboratory of Specialty Fruits and Vegetables Cultivation Physiology and Germplasm Resources Utilization of Xinjiang Production and Construction Corps，Department of Horticulture，Shihezi University，Shihezi，Xinjiang 832000，China

Received:2025-12-23 Revised:2026-03-09 Online:2026-04-25 Published:2026-04-20
Contact: LIU Huiying

Abstract

Abstract:

This study investigated the effects of different inoculation doses of the arbuscular mycorrhizal fungus Funneliformis mosseae on the growth and rhizosphere microenvironment of pepper in biochar-amended sandy soils. The objectives were to determine the optimal inoculation dose of F. mosseae and provide a theoretical and technical basis for improving sand culture substrates and desertified soils for protected-vegetable cultivation in the Gobi Desert of Xinjiang. Pepper（Capsicum annuum‘Lü Xuanfeng'） was used as the experimental plant，and the substrate supplemented with 3%（w/w）rice-husk biochar. The experiment included a non-inoculated control and four F. mosseae doses of 2，4，6，and 8 g · kg^-1 of substrate. The growth parameters，photosynthetic traits and yield of pepper plant，physicochemical properties of the substrate，activities of key enzymes，and rhizosphere microbial community compositions were determined. The results showed that the different inoculation dose of F. mosseae has varying degrees of promoting effects on the growth andyield of sand-cultivated pepper，with 6 g · kg^-1 showing the best application effect. The treatment significantly increased the rate of root colonization，shoot and root dry weights，root morphology and activity. Furthermore，the photosynthetic efficiency and chlorophyll contents of pepper after 6 g · kg^-1 treatment was significantly enhanced. Net photosynthetic rate（P_n）and yield significantly increased by 29.53% and 12.80%，respectively compared with the control. In addition，Analysis of the microbial communities in the rhizosphere soil showed that 6 g · kg^-1 treatment enhanced the relative abundance of beneficial microbial taxa，including Actinomycota，Bacteroidota，Glomeromycota，and Ascomycota，and significantly elevated the contents of available nitrogen，available phosphorus，and available potassium. In conclusion，6 g · kg^-1 was identified as the optimal F. mosseae inoculation dose for promoting pepper growth，increasing fruit yield，and optimizing the rhizosphere microenvironment in biochar-amended sandy soils. The positive effects were attributed primarily to enhanced mycorrhizal symbiosis，improved nutrient uptake by the roots，and increased photosynthesis in the leaves，as well as optimization of nutrient availability in the substrate and microbial community compositions in the rhizosphere soil.

Key words: pepper, sand-cultivated, Funneliformis mosseae, inoculation dose, rhizosphere microecology, growth

YANG Yuqi, WEI Yuxin, LI Xuezhen, KE Xixi, HUANG Tingting, LIU Huiying. Effects of Funneliformis mosseae Inoculation on the Growth and Rhizosphere Microenvironment of Sand-Cultivated Pepper Amended with Biochar[J]. Acta Horticulturae Sinica, 2026, 53(4): 1073-1087.

Figures/Tables 12

References 46

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时间/d Time	接种量/（g · kg^-1） Inoculum doses	株高/cm Height	茎粗/mm Stem diameter	鲜质量/g Fresh weight		干质量/g Dry weight
时间/d Time	接种量/（g · kg^-1） Inoculum doses	株高/cm Height	茎粗/mm Stem diameter	地上部Shoot	地下部Root	地上部Shoot	地下部Root
50	对照Control	14.23 ± 0.41 c	3.01 ± 0.31 b	8.89 ± 0.37 c	2.58 ± 0.09 c	0.27 ± 0.04 b	0.93 ± 0.13 c
	2	14.73 ± 0.39 c	4.02 ± 0.09 a	9.53 ± 0.55 bc	2.77 ± 0.13 bc	0.28 ± 0.05 b	0.95 ± 0.15 bc
	4	17.03 ± 0.56 b	4.18 ± 0.09 a	11.91 ± 1.10 ab	3.46 ± 0.26 ab	0.40 ± 0.06 ab	1.38 ± 0.19 ab
	6	19.80 ± 0.54 a	4.27 ± 0.04 a	13.18 ± 1.22 a	3.62 ± 0.28 a	0.52 ± 0.12 a	1.88 ± 0.44 a
	8	20.77 ± 0.53 a	4.21 ± 0.02 a	12.05 ± 1.02 ab	3.49 ± 0.24 a	0.46 ± 0.12 a	1.45 ± 0.40 ab
60	对照Control	19.80 ± 0.57 c	4.08 ± 0.11 c	15.79 ± 0.76 d	2.92 ± 0.14 d	1.02 ± 0.14 d	0.18 ± 0.03 c
	2	22.50 ± 0.49 b	4.29 ± 0.07 bc	17.90 ± 0.70 cd	3.30 ± 0.12 cd	1.04 ± 0.17 d	0.19 ± 0.04 c
	4	23.73 ± 0.78 b	4.49 ± 0.14 ab	18.81 ± 0.81 bc	3.47 ± 0.14 bc	1.51 ± 0.21 c	0.28 ± 0.04 b
	6	26.13 ± 0.52 a	4.73 ± 0.03 a	21.91 ± 0.70 a	4.04 ± 0.12 a	1.63 ± 0.46 a	0.38 ± 0.09 a
	8	26.07 ± 0.21 a	4.59 ± 0.08 a	21.22 ± 0.40 ab	3.92 ± 0.07 ab	1.61 ± 0.44 b	0.29 ± 0.08 b
70	对照Control	28.40 ± 0.54 c	4.99 ± 0.07 c	26.40 ± 0.79 d	4.46 ± 0.92 b	2.20 ± 0.09 d	0.38 ± 0.02 d
	2	30.20 ± 0.37 c	5.19 ± 0.02 b	28.76 ± 0.67 c	4.57 ± 0.20 b	2.36 ± 0.14 d	0.40 ± 0.03 d
	4	35.73 ± 0.74 b	5.31 ± 0.03 b	33.84 ± 0.40 b	5.39 ± 0.45 ab	2.96 ± 0.28 c	0.50 ± 0.05 c
	6	38.83 ± 0.53 a	5.79 ± 0.04 a	36.09 ± 0.50 a	6.38 ± 0.51 a	3.27 ± 0.29 a	0.55 ± 0.05 a
	8	37.46 ± 0.63 ab	5.75 ± 0.04 a	35.49 ± 0.74 ab	5.54 ± 0.32 ab	2.99 ± 0.26 b	0.51 ± 0.04 b

时间/d Time	接种量/（g · kg^-1） Inoculum doses	株高/cm Height	茎粗/mm Stem diameter	鲜质量/g Fresh weight		干质量/g Dry weight
时间/d Time	接种量/（g · kg^-1） Inoculum doses	株高/cm Height	茎粗/mm Stem diameter	地上部Shoot	地下部Root	地上部Shoot	地下部Root
50	对照Control	14.23 ± 0.41 c	3.01 ± 0.31 b	8.89 ± 0.37 c	2.58 ± 0.09 c	0.27 ± 0.04 b	0.93 ± 0.13 c
	2	14.73 ± 0.39 c	4.02 ± 0.09 a	9.53 ± 0.55 bc	2.77 ± 0.13 bc	0.28 ± 0.05 b	0.95 ± 0.15 bc
	4	17.03 ± 0.56 b	4.18 ± 0.09 a	11.91 ± 1.10 ab	3.46 ± 0.26 ab	0.40 ± 0.06 ab	1.38 ± 0.19 ab
	6	19.80 ± 0.54 a	4.27 ± 0.04 a	13.18 ± 1.22 a	3.62 ± 0.28 a	0.52 ± 0.12 a	1.88 ± 0.44 a
	8	20.77 ± 0.53 a	4.21 ± 0.02 a	12.05 ± 1.02 ab	3.49 ± 0.24 a	0.46 ± 0.12 a	1.45 ± 0.40 ab
60	对照Control	19.80 ± 0.57 c	4.08 ± 0.11 c	15.79 ± 0.76 d	2.92 ± 0.14 d	1.02 ± 0.14 d	0.18 ± 0.03 c
	2	22.50 ± 0.49 b	4.29 ± 0.07 bc	17.90 ± 0.70 cd	3.30 ± 0.12 cd	1.04 ± 0.17 d	0.19 ± 0.04 c
	4	23.73 ± 0.78 b	4.49 ± 0.14 ab	18.81 ± 0.81 bc	3.47 ± 0.14 bc	1.51 ± 0.21 c	0.28 ± 0.04 b
	6	26.13 ± 0.52 a	4.73 ± 0.03 a	21.91 ± 0.70 a	4.04 ± 0.12 a	1.63 ± 0.46 a	0.38 ± 0.09 a
	8	26.07 ± 0.21 a	4.59 ± 0.08 a	21.22 ± 0.40 ab	3.92 ± 0.07 ab	1.61 ± 0.44 b	0.29 ± 0.08 b
70	对照Control	28.40 ± 0.54 c	4.99 ± 0.07 c	26.40 ± 0.79 d	4.46 ± 0.92 b	2.20 ± 0.09 d	0.38 ± 0.02 d
	2	30.20 ± 0.37 c	5.19 ± 0.02 b	28.76 ± 0.67 c	4.57 ± 0.20 b	2.36 ± 0.14 d	0.40 ± 0.03 d
	4	35.73 ± 0.74 b	5.31 ± 0.03 b	33.84 ± 0.40 b	5.39 ± 0.45 ab	2.96 ± 0.28 c	0.50 ± 0.05 c
	6	38.83 ± 0.53 a	5.79 ± 0.04 a	36.09 ± 0.50 a	6.38 ± 0.51 a	3.27 ± 0.29 a	0.55 ± 0.05 a
	8	37.46 ± 0.63 ab	5.75 ± 0.04 a	35.49 ± 0.74 ab	5.54 ± 0.32 ab	2.99 ± 0.26 b	0.51 ± 0.04 b

时间/d Time	接种量/（g · kg^-1） Inoculum doses	总根长/cm Total root length	表面积/cm² Surface area	平均直径/mm Average diameter	根体积/cm³ Root volume	根尖数 Number of root tips
50	对照Control	1 065.98 ± 56.60 d	188.14 ± 13.39 b	0.57 ± 0.02 b	2.90 ± 0.25 d	1 107.00 ± 98.66 c
	2	1 259.81 ± 37.98 c	245.77 ± 20.14 ab	0.64 ± 0.03 ab	4.08 ± 0.23 c	1 456.67 ± 84.53 b
	4	1 361.14 ± 68.72 bc	275.58 ± 34.28 a	0.65 ± 0.03 ab	4.90 ± 0.65 bc	1 781.67 ± 56.88 a
	6	1 556.77 ± 52.16 a	307.49 ± 31.29 a	0.70 ± 0.06 a	6.61 ± 0.17 a	1 812.67 ± 39.72 a
	8	1 506.81 ± 55.32 ab	311.34 ± 25.14 a	0.69 ± 0.02 ab	5.41 ± 0.20 b	1 580.67 ± 67.06 b
60	对照Control	1 268.00 ± 26.92 c	244.85 ± 14.86 b	0.72 ± 0.02 c	4.67 ± 0.25 b	1 040.00 ± 42.09 c
	2	1 354.14 ± 70.35 bc	305.70 ± 47.98 ab	0.74 ± 0.02 c	5.52 ± 0.27 b	1 306.67 ± 44.32 bc
	4	1 441.62 ± 53.03 bc	338.51 ± 26.13 ab	0.76 ± 0.02 bc	7.16 ± 0.20 a	1 549.33 ± 43.28 ab
	6	1 684.78 ± 66.94 a	377.92 ± 45.74 a	0.83 ± 0.01 a	7.40 ± 0.32 a	1 841.67 ± 77.37 a
	8	1 558.25 ± 59.24 ab	373.54 ± 33.12 a	0.84 ± 0.05 a	7.24 ± 0.38 a	1 580.33 ± 34.72 ab
70	对照Control	1 319.00 ± 47.43 d	366.81 ± 25.65 b	0.84 ± 0.02 b	7.73 ± 0.56 c	1 114.00 ± 26.94 c
	2	1 485.22 ± 46.31 c	431.12 ± 20.95 ab	0.91 ± 0.02 b	11.04 ± 0.71 b	1 396.33 ± 73.74 bc
	4	1 561.17 ± 33.33 bc	444.71 ± 50.07 ab	0.92 ± 0.03 b	12.36 ± 0.58 ab	1 401.33 ± 39.69 bc
	6	1 768.92 ± 53.33 a	475.13 ± 46.80 ab	1.09 ± 0.04 a	14.48 ± 0.80 a	1 766.67 ± 73.41 a
	8	1 650.21 ± 17.60 ab	532.66 ± 31.04 a	0.94 ± 0.03 b	13.16 ± 0.59 ab	1 503.67 ± 39.20 ab

时间/d Time	接种量/（g · kg^-1） Inoculum doses	总根长/cm Total root length	表面积/cm² Surface area	平均直径/mm Average diameter	根体积/cm³ Root volume	根尖数 Number of root tips
50	对照Control	1 065.98 ± 56.60 d	188.14 ± 13.39 b	0.57 ± 0.02 b	2.90 ± 0.25 d	1 107.00 ± 98.66 c
	2	1 259.81 ± 37.98 c	245.77 ± 20.14 ab	0.64 ± 0.03 ab	4.08 ± 0.23 c	1 456.67 ± 84.53 b
	4	1 361.14 ± 68.72 bc	275.58 ± 34.28 a	0.65 ± 0.03 ab	4.90 ± 0.65 bc	1 781.67 ± 56.88 a
	6	1 556.77 ± 52.16 a	307.49 ± 31.29 a	0.70 ± 0.06 a	6.61 ± 0.17 a	1 812.67 ± 39.72 a
	8	1 506.81 ± 55.32 ab	311.34 ± 25.14 a	0.69 ± 0.02 ab	5.41 ± 0.20 b	1 580.67 ± 67.06 b
60	对照Control	1 268.00 ± 26.92 c	244.85 ± 14.86 b	0.72 ± 0.02 c	4.67 ± 0.25 b	1 040.00 ± 42.09 c
	2	1 354.14 ± 70.35 bc	305.70 ± 47.98 ab	0.74 ± 0.02 c	5.52 ± 0.27 b	1 306.67 ± 44.32 bc
	4	1 441.62 ± 53.03 bc	338.51 ± 26.13 ab	0.76 ± 0.02 bc	7.16 ± 0.20 a	1 549.33 ± 43.28 ab
	6	1 684.78 ± 66.94 a	377.92 ± 45.74 a	0.83 ± 0.01 a	7.40 ± 0.32 a	1 841.67 ± 77.37 a
	8	1 558.25 ± 59.24 ab	373.54 ± 33.12 a	0.84 ± 0.05 a	7.24 ± 0.38 a	1 580.33 ± 34.72 ab
70	对照Control	1 319.00 ± 47.43 d	366.81 ± 25.65 b	0.84 ± 0.02 b	7.73 ± 0.56 c	1 114.00 ± 26.94 c
	2	1 485.22 ± 46.31 c	431.12 ± 20.95 ab	0.91 ± 0.02 b	11.04 ± 0.71 b	1 396.33 ± 73.74 bc
	4	1 561.17 ± 33.33 bc	444.71 ± 50.07 ab	0.92 ± 0.03 b	12.36 ± 0.58 ab	1 401.33 ± 39.69 bc
	6	1 768.92 ± 53.33 a	475.13 ± 46.80 ab	1.09 ± 0.04 a	14.48 ± 0.80 a	1 766.67 ± 73.41 a
	8	1 650.21 ± 17.60 ab	532.66 ± 31.04 a	0.94 ± 0.03 b	13.16 ± 0.59 ab	1 503.67 ± 39.20 ab

时间/d Time	接种量/（g · kg^-1） Inoculum doses	叶绿素含量/ （mg · g^-1） Chl content	胞间CO₂浓度/ （µmol · mol^-1） C_i	净光合速率/ (µmol · m^-2 · s^-1) P_n	蒸腾速率/ (mol · m^-2 · s^-1) T_r	气孔导度/ (mol · m^-2 · s^-1) G_s
50	对照Control	2.73 ± 0.10 c	213.15 ± 1.12 b	9.53 ± 0.86 c	4.18 ± 0.29 c	0.37 ± 0.05 a
	2	2.88 ± 0.08 bc	210.39 ± 5.55 b	10.45 ± 0.35 bc	4.72 ± 0.30 c	0.40 ± 0.04 a
	4	2.97 ± 0.05 ab	257.49 ± 4.65 a	10.52 ± 0.42 bc	5.10 ± 0.46 bc	0.33 ± 0.01 a
	6	3.16 ± 0.04 a	266.31 ± 3.12 a	13.09 ± 0.67 a	6.04 ± 0.34 a	0.30 ± 0.02 a
	8	3.05 ± 0.09 ab	260.33 ± 3.43 a	11.20 ± 0.40 b	5.91 ± 0.41 ab	0.29 ± 0.03 a
60	对照Control	2.82 ± 0.05 d	201.82 ± 8.49 c	10.14 ± 0.37 d	5.86 ± 0.80 b	0.46 ± 0.01 a
	2	3.00 ± 0.06 c	208.65 ± 2.77 c	12.24 ± 0.29 c	7.40 ± 0.65 ab	0.35 ± 0.01 b
	4	3.14 ± 0.01 b	237.96 ± 5.35 b	12.05 ± 0.25 c	7.55 ± 1.07 ab	0.32 ± 0.02 b
	6	3.28 ± 0.05 a	259.73 ± 3.98 a	14.58 ± 0.08 a	8.32 ± 0.03 a	0.25 ± 0.01 c
	8	3.30 ± 0.01 a	257.36 ± 4.04 a	13.07 ± 0.23 b	7.40 ± 0.35 ab	0.26 ± 0.02 c
70	对照Control	2.76 ± 0.03 c	180.99 ± 4.60 d	10.84 ± 0.39 c	4.88 ± 0.06 c	0.46 ± 0.02 a
	2	3.03 ± 0.02 b	195.77 ± 4.44 c	13.80 ± 0.25 b	6.38 ± 0.21 b	0.40 ± 0.02 b
	4	3.04 ± 0.02 b	214.97 ± 4.22 b	14.74 ± 0.60 ab	6.96 ± 0.31 b	0.36 ± 0.01 c
	6	3.15 ± 0.02 a	231.46 ± 2.98 a	15.41 ± 0.55 a	8.27 ± 0.37 a	0.32 ± 0.01 c
	8	3.19 ± 0.06 a	224.70 ± 5.07 ab	14.81 ± 0.56 ab	7.68 ± 0.64 a	0.32 ± 0.01 c

Effects of Funneliformis mosseae Inoculation on the Growth and Rhizosphere Microenvironment of Sand-Cultivated Pepper Amended with Biochar

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接种量/ （g · kg^-1） Inoculum doses	容重/（g · cm^-3） Bulk density	总孔隙度/% Total porosity	通气孔隙/% Aeration porosity	持水孔隙/% Water-holding porosity	大小孔隙比 Ratio of aeration porosity to water-holding porosity	pH
对照Control	1.02 ± 0.01 a	50.53 ± 0.40 b	41.26 ± 0.47 a	8.79 ± 0.60 a	0.21 ± 1.28 a	8.597 ± 0.012 b
2	1.01 ± 0.01 a	51.10 ± 0.16 ab	41.79 ± 0.59 a	9.31 ± 0.72 a	0.22 ± 1.22 a	8.593 ± 0.005 b
4	1.02 ± 0.01 a	51.11 ± 0.35 ab	41.83 ± 0.72 a	9.28 ± 0.53 a	0.22 ± 0.74 a	8.623 ± 0.021 b
6	1.01 ± 0.01 a	51.39 ± 0.08 a	41.86 ± 0.16 a	9.20 ± 0.52 a	0.22 ± 3.22 a	8.590 ± 0.016 b
8	1.01 ± 0.02 a	51.24 ± 0.69 ab	41.78 ± 1.02 a	9.46 ± 0.79 a	0.23 ± 0.77 a	8.680 ± 0.016 a

接种量/ （g · kg^-1） Inoculum doses	有机质/ （g · kg^-1） Organic matter	全氮/ （g · kg^-1） Total nitrogen	碱解氮/ （mg · kg^-1） Alkali-hydroly zable nitrogen	全磷/ （g · kg^-1） Total phosphorus	有效磷/ （mg · kg^-1） Available phosphorus	全钾/ （g · kg^-1） Total potassium	速效钾/ （mg · kg^-1） Available potassium
对照Control	6.09 ± 0.05 b	0.16 ± 0.01 b	10.76 ± 0.13 c	0.50 ± 0.01 a	13.13 ± 0.12 d	21.74 ± 0.14 a	677.97 ± 0.69 c
2	6.11 ± 0.02 b	0.18 ± 0.01 b	11.52 ± 0.49 bc	0.48 ± 0.01 b	15.62 ± 0.36 c	22.00 ± 0.24 a	738.44 ± 2.98 b
4	6.21 ± 0.06 b	0.16 ± 0.01 b	13.04 ± 0.36 ab	0.47 ± 0.01 b	15.15 ± 0.47 c	22.12 ± 0.04 a	742.00 ± 6.07 b
6	6.65 ± 0.11 a	0.23 ± 0.02 a	14.85 ± 1.28 a	0.47 ± 0.01 b	19.12 ± 0.27 a	22.11 ± 0.65 a	857.79 ± 0.48 a
8	6.55 ± 0.11 a	0.19 ± 0.01 b	12.85 ± 0.47 abc	0.48 ± 0.01 b	17.00 ± 0.39 b	22.09 ± 0.15 a	848.87 ± 1.75 a

接种量/（g · kg^-1） Inoculum doses	碱性磷酸酶/ （μmol · g^-1 · d^-1） Alkaline phosphatase	酸性磷酸酶/ （μmol · g^-1 · d^-1） Acid phosphatase	脲酶/ （μg · g^-1 · d^-1） Urease	脱氢酶/ （(μg · g^-1 · d^-1） Dehydrogenase	纤维二糖酶/ （μmol · g^-1 · d^-1） Cellobiosidase
对照Control	1.15 ± 0.08 c	1.47 ± 0.02 c	175.70 ± 5.06 d	15.95 ± 0.65 c	5.28 ± 0.77 b
2	1.22 ± 0.05 bc	1.74 ± 0.05 bc	193.18 ± 4.80 c	18.68 ± 0.79 c	6.55 ± 0.67 ab
4	1.43 ± 0.09 ab	2.54 ± 0.31 a	221.44 ± 3.81 b	35.07 ± 0.21 b	7.02 ± 0.62 ab
6	1.55 ± 0.08 a	2.60 ± 0.36 a	256.11 ± 5.03 a	47.31 ± 1.02 a	7.92 ± 0.69 a
8	1.47 ± 0.05 ab	2.50 ± 0.10 ab	251.16 ± 6.62 a	36.65 ± 1.50 b	6.04 ± 0.49 ab

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