Chin J Plan Ecolo ›› 2015, Vol. 39 ›› Issue (12): 1198-1208.doi: 10.17521/cjpe.2015.0116

• Orginal Article • Previous Articles     Next Articles

Interactive effects of phenolic acid and nitrogen on morphological traits of poplar (Populus × euramericana ‘Neva’) fine roots

ZHU Wan-Rui1, WANG Qi-Tong1, LIU Meng-Ling1, WANG Hua-Tian1,2, WANG Yan-Ping1,2,*(), ZHANG Guang-Can1,2, LI Chuan-Rong1,2   

  1. 1Forestry College of Shandong Agricultural University, Tai’an, Shandong 271018, China
    and 2Taishan Forest Ecosystem Research Station of State Forestry Administration, Tai’an, Shandong 271018, China
  • Online:2015-12-31 Published:2015-12-01
  • Contact: Yan-Ping WANG
  • About author:

    # Co-first authors


Aims The relationship between rhizosphere process and fine root growth is very close but still obscure. In poplar plantation, phenolic acid rhizodeposition and soil nutrient availability were considered as two dominant factors of forest productivity decline. It is very hard to separate them in the field and they might show an interactive effect on fine root growth. The objective of this study is to examine the influence of phenolic acids and nitrogen on branch orders of poplar fine roots and to give a deeper insight into how the ecological process on root-soil interface affected fine root growth as well as plantation productivity. Methods The cuttings of health annual poplar seedlings (I-107, Populus × euramericana ‘Neva’) serve as experiment materials, and were cultivated under nine conditions, including three concentration of phenolic acids at 0X, 0.5X, 1.0X (here, X represented the contents of phenolic acids in the soil of poplar plantation) and three concentration of nitrogen at 0 mmol·L-1, 10 mmol·L-1, 20 mmol·L-1, based on Hoagland solution. The roots were all separated from poplar seedlings after 35 days, and 30 percent of total fine roots of every treatment were taken as fine root samples. These fine roots were grouped according to 1 to 5 branch orders, and then the morphological traits of each group of fine roots were scanned via root analyzer system (WinRHIZO, Regent Instruments Company, Quebec, Canada) including total length, surface area, volume and average diameter. Meanwhile, the dry mass of fine root samples of every order was measured to calculate specific root length (SRL), root tissue density (RTD). All data were analyzed via SPSS 17.0 software, and interactive effect of phenolic acids and nitrogen on roots was analyzed through univariate process module. Principal component analysis (PCA) and redundancy analysis (RDA) were conducted via Canoco 4.5 software. Important findings Under the conditions without phenolic acids application, the fine roots growth was significantly inhibited in deficiency and higher nitrogen treatments, especially for 1-3 order roots. Only specific root length appeared decreased with nitrogen level, and other traits of fine roots did not demonstrate linear relationship with nitrogen concentrations. Compared to 0.5X phenolic acids treatment, 1.0X phenolic acids significantly promoted the diameter and volume of 1-2 order roots (p < 0.05). Both phenolic acids and nitrogen demonstrated influence on poplar fine root traits. However, the diameter and volume of 1-2 order roots were significantly affected by phenolic acids, while the total length and surface area of 4-5 order roots was affected by nitrogen. Two way ANOVA showed that phenolic acids and nitrogen made a synergistic or antagonistic effect on morphological building of fine roots. Furthermore, PCA and RDA indicated that the interactive effects of phenolic acids and nitrogen led to significant differences among 1-3 order, 4th order and 5th order of poplar fine roots. The PC1 explained about 60.9 percent of root morphological variance, which was related to foraging traits of roots. The PC2 explained 25.3 percent of variance, which was related to root building properties. The response of poplar roots to phenolic acids and nitrogen was closely related to root order, and nitrogen played more influence on poplar roots than phenolic acids. Thus, phenolic acids and nitrogen level would affect many properties of root morphology and foraging in rhizosphere soil of poplar plantation. But nitrogen availability would serve as a dominant factor influencing root growth, and soil nutrient management should be critical to productivity maintenance of poplar plantation.

Key words: fine root morphological traits, nitrogen availability, phenolic acids rhizodeposition, poplar root order, root-soil interaction

Table 1

Phenolic acid concentration setting in the treatment solutions"

p-hydroxybenzoic acid
Ferulic acid
Benzoic acid
Cinnamic acid
Content of phenolic acid in field soil (μg·g-1)
152.00 10.40 6.50 20.60 1.95
Absorption rate of phenolic acid in soil (%)
61.66 93.78 89.30 37.85 94.41
Gradient of phenolic acid content (μg·mL-1)
0.5X 123 6 4 27 1
1.0X 247 11 7 54 2

Table 2

The response of morphological traits of different order fine roots to the nitrogen treatments"

根序 Root order 氮素水平
Nitrogen level
Total root
Total root
surface area
Total root
Root dry
Specific root
Root tissue
1 0 534.51 ± 58.78a 28.93 ± 2.11a 0.13 ± 0.005a 0.17 ± 0.064a 0.013 ± 0.000a 323.00 ± 12.66a 0.114 ± 0.010ab
10 822.03 ± 65.15b 55.41 ± 9.47b 0.41 ± 0.007b 0.23 ± 0.013b 0.028 ± 0.002b 287.53 ± 4.28b 0.091 ± 0.015a
20 389.77 ± 48.70a 23.55 ± 2.28a 0.12 ± 0.011a 0.18 ± 0.025a 0.014 ± 0.002a 246.51 ± 37.71b 0.146 ± 0.001b
2 0 716.81 ± 49.22a 42.01 ± 2.90a 0.19 ± 0.013a 0.19 ± 0.001a 0.023 ± 0.003a 269.07 ± 36.03a 0.113 ± 0.003a
10 871.82 ± 15.89a 72.59 ± 3.10b 0.26 ± 0.017b 0.25 ± 0.011b 0.042 ± 0.002b 228.89 ± 18.80ab 0.085 ± 0.001b
20 428.53 ± 61.70b 39.30 ± 1.34a 0.23 ± 0.008ab 0.21 ± 0.006a 0.023 ± 0.001a 185.39 ± 10.71b 0.108 ± 0.001a
3 0 249.26 ± 19.75a 16.37 ± 0.22a 0.10 ± 0.007a 0.24 ± 0.016a 0.013 ± 0.001a 184.68 ± 22.25a 0.104 ± 0.004a
10 462.39 ± 52.25b 28.30 ± 3.85b 0.22 ± 0.014b 0.30 ± 0.037a 0.026 ± 0.000b 146.59 ± 11.16ab 0.118 ± 0.006ab
20 179.04 ± 16.34a 16.42 ± 1.27a 0.14 ± 0.026a 0.29 ± 0.003a 0.017 ± 0.002a 111.33 ± 2.83b 0.129 ± 0.005b
4 0 38.28 ± 4.84a 4.39 ± 0.30a 0.06 ± 0.002a 0.32 ± 0.016a 0.006 ± 0.000a 75.49 ± 7.93a 0.108 ± 0.002a
10 43.26 ± 4.42b 5.62 ± 0.47ab 0.07 ± 0.009b 0.42 ± 0.037a 0.008 ± 0.003a 63.54 ± 5.19b 0.122 ± 0.005a
20 39.29 ± 4.19b 4.52 ± 0.91b 0.04 ± 0.011a 0.39 ± 0.016a 0.005 ± 0.002a 60.92 ± 1.38b 0.130 ± 0.003a
5 0 32.85 ± 1.14a 7.41 ± 0.44a 0.17 ± 0.024a 0.74 ± 0.018a 0.032 ± 0.001a 12.58 ± 1.83a 0.155 ± 0.004a
10 48.15 ± 4.71a 13.85 ± 1.63a 0.47 ± 0.046a 1.03 ± 0.056a 0.043 ± 0.003a 9.85 ± 1.06b 0.114 ± 0.284a
20 28.24 ± 11.61a 7.74 ± 4.25a 0.17 ± 0.118a 0.74 ± 0.136a 0.024 ± 0.016a 8.69 ± 0.49b 0.140 ± 0.003a

Fig. 1

The fine root morphological characteristics in the interaction of phenolic acid and nitrogen treatments (mean ± SE). Different lowercase letters and capital letters represent significant difference among three nitrogen treatments under 0.5X and 1.0X phenolic acids respectively (p < 0.05), here X represents the contents of phenolic acids in the soil of poplar plantation. Three symbols, +, * and #, represent significant difference between two treatments of phenolic acids under nitrogen deficiency (0 mmol·L-1), normal nitrogen (10 mmol·L-1) and high nitrogen (20 mmol·L-1), respectively."

Fig. 2

The dry mass, specific root length and root tissue density of different fine root orders under the interaction of phenolic acid and nitrogen conditions (mean ± SE). Different lowercase letters and capital letters represent significant difference among three nitrogen treatments under 0.5X and 1.0X phenolic acids respectively (p < 0.05), here X represents the contents of phenolic acids in the soil of poplar plantation. Three symbols, +, * and #, represent significant difference between two treatments of phenolic acids under nitrogen deficiency (0 mmol·L-1), normal nitrogen (10 mmol·L-1) and high nitrogen (20 mmol·L-1), respectively."

Table 3

The interaction effect of phenolic acid and nitrogen on morphological traits of poplar fine roots"

根序 Root order 处理
Total root
Length (cm)
Total root surface area (cm2)
Total root
volume (cm3)
Average diameter
of root (mm)
Root dry
mass (g)
Specific root length (m·g-1)
Root tissue density (g·cm-3)
First order
pN 0.002* 0.000* 0.000* 0.000* 0.009* 0.018* 0.001*
pT 0.116ns 0.559ns 0.295ns 0.201ns 0.023* 0.048* 0.667ns
p(N×T) 0.300ns 0.394ns 0.003* 0.867ns 0.067ns 0.852ns 0.008*
Second order
pN 0.005* 0.015* 0.071ns 0.001* 0.001* 0.000* 0.009*
pT 0.041* 0.054ns 0.569ns 0.083ns 0.002* 0.715ns 0.164ns
p(N×T) 0.015* 0.028* 0.557ns 0.424ns 0.001* 0.853ns 0.011*
Third order
pN 0.039* 0.014* 0.080ns 0.003* 0.017* 0.000* 0.000*
pT 0.481ns 0.011* 0.046* 0.164ns 0.035* 0.366ns 0.931ns
p(N×T) 0.004* 0.012* 0.086ns 0.923ns 0.118ns 0.494ns 0.023*
Forth order
pN 0.001* 0.021* 0.503ns 0.001* 0.097ns 0.000* 0.052ns
pT 0.066ns 0.764ns 0.406ns 0.066ns 0.379ns 0.409ns 0.521ns
p(N×T) 0.220ns 0.926ns 0.070ns 0.220ns 0.121ns 0.634ns 0.819ns
Fifth order
pN 0.005* 0.048* 0.012* 0.067ns 0.108ns 0.002* 0.222ns
pT 0.631ns 0.238ns 0.111ns 0.453ns 0.163ns 0.092ns 0.876ns
p(N×T) 0.760ns 0.824ns 0.552ns 0.129ns 0.884ns 0.124ns 0.569ns

Table 4

Component matrix of fine root growth indices"

Morphological traits of fine root
主成分 Principal component
1 2
根长 Total root length 0.872 0.421
根表面积 Total root surface area 0.912 0.542
根体积 Total root volume 0.239 0.916
根直径 Average diameter of root -0.805 0.246
根干质量 Root dry mass 0.000 0.952
比根长 Specific root length 0.956 0.069
根组织密度 Root tissue density -0.655 -0.330


Fig. 4

Redundancy analysis (RDA) on the effects of phenolic acids and nitrogen on morphological traits of poplar fine roots. Real vectors represent environmental factors (phenolic acids and nitrogen), and dotted vectors represent morphological traits of poplar fine roots (the same indices with Table 2). The longer vector is the more important the environmental effects. The correlation between the variables is illustrated by the angle between two vectors. Vectors pointing in nearly the same direction indicate a high positive correlation, vectors pointing in opposite directions have a high negative correlation, and vectors crossing at right angles are related to a near zero correlation."

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