Chin J Plan Ecolo ›› 2015, Vol. 39 ›› Issue (1): 32-42.doi: 10.17521/cjpe.2015.0004

• Orginal Article • Previous Articles     Next Articles

Responses of the functional traits in Cleistogenes squarrosa to nitrogen addition and drought

YANG Hao1,*(), LUO Ya-Chen1,2   

  1. 1Key Laboratory of Ecosystem Network Observation and Modeling, Institute of Geographical Sciences and Natural Resources Research, Chinese Academy of Sciences, Beijing 100101, China
    and 2School of Geographical Sciences, Southwest University, Chongqing 400715, China
  • Received:2014-08-04 Accepted:2014-12-09 Online:2015-01-22 Published:2015-01-10
  • Contact: Hao YANG
  • About author:

    # Co-first authors

Abstract: Aims

Plant functional traits have been widely used to study the responses of plant to environmental change. Cleistogenes squarrosa is an important C4 species in Nei Mongol grassland. How its functional traits would respond to varied nitrogen and water conditions have rarely been studied. Our subject was to examine the responses of the whole-plant traits and leaf morphological and physiological traits to nitrogen addition and drought in this species.


We conducted a pot experiment with a gradient of N addition (0, 10.5, 35.0, and 56.0 g·m-2·a-1) and water treatments (natural precipitation vs. 70% of mean monthly precipitation) in 2013. The whole-plant traits (e.g., root depth and stem-leaf biomass ratio), leaf morphological (e.g., leaf area and specific leaf area) and physiological traits (e.g., photosynthetic rate, water use efficiency, and leaf N content) were investigated.

<i>Important findings </i>

N addition had a significant effect on the whole-plant traits in C. squarrosa. The effects of N addition, water treatments, and an interaction between N addition and water treatments on leaf morphological and physiological traits were highly significant in most cases. The patterns of functional traits in response to N addition differed between plants under natural precipitation and with reduced mean monthly precipitation. Root depth, stem biomass, and stem-leaf biomass ratio were increased in treatments with low and intermediate N additions under reduced precipitation, but not changed under natural precipitation. Specific leaf area increased along the N addition gradient under drought, but did not change under natural precipitation. High N addition stimulated photosynthetic rate and transpiration rate and increased water use efficiency under natural precipitation, but had no effect on under reduced precipitation. Leaf N content on area basis increased slightly with the increases in N addition under natural precipitation, but decreased significantly under reduced precipitation. N addition influenced mainly the leaf morphological and physiological traits under natural precipitation and the whole-plant traits and leaf morphological traits under reduced precipitation. In conclusion, our results indicate that the functional traits in C. squarrosa respond to N addition and the patterns of responses differ under different water conditions, reflecting the adaptation to changes in N and water availability.

Key words: C4 plant, leaf nitrogen content, morphological traits, Nei Mongol grassland, photosynthetic rate, physiological traits

Fig. 1

Plot layout."

Table 1

Results of two-way ANOVAs for the effects of N addition, water treatments and their interaction on whole-plant traits and leaf morphological traits in Cleistogenes squarrosa"

整株性状 Whole-plant traits 叶形态性状 Leaf morphological traits
Height (cm)
Root depth (cm)
Stem biomass (mg·tiller-1)
Leaf biomass (mg·tiller-1)
Stem: leaf ratio (tiller-1)
Leaf area (cm2·leaf-1)
Leaf dry mass (mg·leaf-1)
Specific leaf area (cm2·g-1·leaf-1)
F p F p F p F p F p F p F p F p
N addition
4.930 0.010* 15.340 0.000** 16.940 0.000** 6.080 0.004** 12.040 0.000** 2.940 0.059 0.000 0.986 3.540 0.035*
Water treatment
0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 2.870 0.106 3.680 0.031* 5.480 0.030*
N addition × Water treatment
0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 0.000 1.000 5.950 0.005** 8.030 0.001** 5.310 0.008**

Fig. 2

Effects of N addition and drought on whole-plant traits in Cleistogenes squarrosa (mean ± SE, n = 4). Difference letters indicate significant differences among treatments (LSD test, p < 0.05)."

Table 2

Results of two-way ANOVAs for the effects of N addition, water treatments and their interaction on leaf photosynthetic traits and leaf nitrogen content in Cleistogenes squarrosa"

叶生理形状 Leaf photosynthetic traits 叶片氮含量 Leaf nitrogen content
(μmol CO2·m-2·s-1)
(mmol H2O·m-2·s-1)
(mol H2O·m-2·s-1)
(μmol CO2·mol-1)
(μmol CO2·mol-1 H2O)
Nmass (%)
Narea (g·m-2)
F p F p F p F p F p F p F p F p
N addition
3.91 0.024* 1.86 0.169 1.39 0.274 11.28 0.000** 2.44 0.095 11.49 0.000** 6.71 0.003** 1.34 0.290
Water treatment
9.19 0.007** 12.89 0.002** 10.85 0.004** 6.29 0.021* 0.44 0.513 6.50 0.019* 7.55 0.012* 15.05 0.001**
N addition × Water treatment
7.82 0.001** 2.70 0.073 4.14 0.020* 10.03 0.000** 3.71 0.029* 9.91 0.000** 0.45 0.720 5.16 0.008**

Fig. 3

Effects of N addition and drought on leaf morphological traits in Cleistogenes squarrosa (mean ± SE, n = 4). Difference letters indicate significant differences among treatments (LSD test, p < 0.05)."

Fig. 4

Effects of N addition and drought on leaf photosynthetic traits in Cleistogenes squarrosa (mean ± SE, n = 4). Ci, intercellular CO2 concentration; Gs, stomatal conductance; Ls, stomatal limitation; Pn, net photosynthetic rate; Tr, transpiration rate; WUE, water use efficiency. Difference letters indicate significant differences among treatments (LSD test, p < 0.05)."

Fig. 5

Effects of N addition and drought on leaf nitrogen content based on leaf mass (Nmass) and leaf nitrogen content based on leaf area (Narea) in Cleistogenes squarrosa (mean ± SE, n = 4). Difference letters indicate significant differences among treatments (LSD test, p < 0.05)."

Fig. 6

The cluster dendrogram of plant traits. Ci, intercellular CO2 concentration; Gs, stomatal conductance; Ls, stomatal limitation; Narea, leaf nitrogen content based on leaf area; Nmass, leaf nitrogen content based on leaf mass; Pn, net photosynthetic rate; Tr, transpiration rate; WUE, water use efficiency."

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