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分根区灌溉和氮肥处理对星油藤幼苗的影响(英文)



全 文 :Effects of Alternative Partial Root-zone Irrigation
and Nitrogen Fertilizer on Plukenetia volubilis
Seedlings
Yanjing GENG1,2, Chuantao CAI1, Zhiquan CAI1*
1. Key Laboratory of Tropical Plant Resources and Sustainable Use, Xishuangbanna Tropical Botanical Garden, Chinese Acade-
my of Sciences, Mengla 666303, China;
2. University of Chinese Academy of Sciences, Beijing 100049, China
Supported by National Natural Science Foundation of China (31370684); Knowledge
Innovation Project of the Chinese Academy of Sciences (KSCX2EWQ17,
KSCX2EWQ15).
*Corresponding author. E-mail: zhiquan.cai@126.com
Received: January 14, 2016 Accepted: March 16, 2016A
Agricultural Science & Technology, 2016, 17(4): 890-895
Copyright訫 2016, Information Institute of HAAS. All rights reserved Soil and Fertilizer
P lukenetia volubilis L. (Euphor-biaceae) is a perennial woodyvine. The seeds of P. volubilis
are rich in fat (45% -60% ), protein
(27%-33%), vitamins, sterols and oth-
er biologically active substances. Am-
ong multi-unsaturated fatty acid,
Omega fatty acid is the dominant com-
ponent[1-2].The oil of P. volubilis is con-
sidered one of the worlds best edible
vegetable oils[3-5]. P. volubilis was intro-
duced to Xishuangbanna Tropical
Botanical Garden in 2006. Nowadays,
it has a certain scale of planting in
Xishuangbanna and Laos.
Due to monsoon climate, annual
rainfall distribution is usually uneven in
tropical and subtropical regions with a
long dry season (November to April
next year). In the dry season, plant
growth is inevitably affected by water
stress. Previous studies have shown
that compared with those under full ir-
rigation, the female flowers, male
flowers, fruits and source-sink ratio of
P. volubilis plants under natural
drought stress were significantly re-
duced, and the abortion rate of fruits
was increased in Xishuangbanna, a
tropical region in southwest China [6].
The developed fruits in the late dry
season are probably to be aborted
when they enter rainy season, result-
ing in low fruit yield in early rainy sea-
son [6]. Therefore, dry-season irrigation
is necessary for the yield improvement
of P. volubilis plants. Southwest China
is short of water resources, especially
in the last several years when serious
drought appeared. It’s an inevitable
Abstract This study was aimed to investigate the effects of alternative partial root-
zone irrigation and nitrogen fertilizer on the potted seedlings of Plukenetia volubilis.
A total of 7 treatments were designed with three factors, i.e., irrigation amount, irri-
gation mode and nitrogen fertilizer. The growth, photosynthesis and water use effi-
ciency were analyzed. The results showed that compared with those under full irri-
gation, the biomass and water consumption under alternative partial root-zone irri-
gation were reduced by 5% and 75% , respectively, and the water use efficiency
was increased by 60%. Under severe drought conditions, the root cap ratio in the
nitrogen fertilizer treatment group was increased by 30%; the leaf area index, photo-
synthetic rate and biomass under alternative partial root-zone irrigation were reduced
by 38%, 9% and 18%, respectively. It indicates that under severe drought condi-
tions, alternative partial root-zone irrigation is not suitable to be matched with appli-
cation of nitrogen fertilizer. In short, under moderate drought conditions, alternative
partial root-zone irrigation could reduce transpiration and improve water use efficien-
cy, and it is an effective water-saving irrigation technology for the plantation of P.
volubilis plants.
Key words Plukenetia volubilis L.; Alternative partial root-zone irrigation; Nitrogen
fertilizer; Growth; Water-use efficiency
分根区灌溉和氮肥处理对星油
藤幼苗的影响
耿艳菁 1,2,蔡传涛 1,蔡志全 1* (1.中国科学院
西双版纳热带植物园热带植物资源可持续利
用重点实验室,云南勐腊 666303;2.中国科学
院大学,北京 100049)
摘 要 探讨分根区灌溉和氮肥对盆栽星油
藤幼苗生长的影响。 试验设置 3 个因素,灌水
量、灌溉方式和氮肥,共 7 个处理,测量了幼苗
生长、光合和水分利用效率等指标。 结果表明:
与充分灌溉相比 ,分根区灌溉的处理,生物量
降低 5%,节水 75%,水分利用率高出 60%,表
现出极大的节水效益。 重度干旱条件下,施氮
肥的处理,根冠比增加达 30%,但分根区灌溉
处理叶面积指数下降 38%,光合速率下降 9%,
生物量下降 18%,分根区灌溉在严重干旱条件
不适宜施氮肥。 总之,在适度干旱条件下,分根
区灌溉降低蒸腾 ,提高水分利用率 ,是一种有
效的节水灌溉技术。
关键词 星油藤;分根区灌溉;氮肥;水分利用
效率
基 金 项 目 国 家 自 然 科 学 基 金 项 目
(31370684);中国科学院知识创新工程重大部
署项目(KSCX2EWQ17,KSCX2EWQ15)。
作者简介 耿艳菁(1987-),女,河南商丘人,硕
士,研究方向:植物生理学。 *通讯作者,E-mail:
zhiquan.cai@126.com。
收稿日期 2016-01-14
修回日期 2016-03-16
DOI:10.16175/j.cnki.1009-4229.2016.04.030
Agricultural Science & Technology2016
Table 1 Irrigation treatments of P. volubilis seedlings
Treatment//% Times Irrigation mode Fertilization Total irrigationamount//ml
NS 26 Full irrigation - 3 372
MD 26 Full irrigation - 1 680
MD+RD 26 Alternative partial root-zone irrigation - 1 315
SD 26 Full irrigation - 684
SD+N 26 Full irrigation √ 695
SD+RD 26 Alternative partial root-zone irrigation - 350
SD+RD+N 26 Alternative partial root-zone irrigation √ 325
NS, normal supply, the irrigation amount is the same with the transpiration; MD, mild
drought, the irrigation amount is 75% of the transpiration; MD +RD, mild drought and
alternative partial root-zone irrigation (RD); SD, severe drought, the irrigation amount is
50% of the transpiration; SD+N, severe drought and N fertilizer (N); SD +RD, severe
drought and alternative partial root-zone irrigation; SD+RD+N, severe drought, alternative
partial root-zone irrigation and N fertilizer.
development tendency for modern a-
griculture to improve water use effi-
ciency and yield with developed water-
saving irrigation technologies.
Alternative partial root-zone irri-
gation (partial root-zone drying) is a
new water-saving technology, which
has attracted much attention [7]. Based
on the theory of root-sourced signaling,
alternative partial root-zone irrigation
could control dried roots, so it has been
widely promoted and applied in maize,
soybeans and other field crops, and
grapes and other fruit trees[8-12].Howev-
er, there has been no report on appli-
cation of alternative partial root-zone ir-
rigation in P. volubilis plants. Although
alternative partial root-zone irrigation
has been widely used in arid and semi-
arid regions, there are few researches
in tropical and subtropical regions [13].
Based on indoor pot experiment, the
effects of alternative partial root-zone
irrigation on growth and physiology of
young P. volubilis seedlings were in-
vestigated in this paper, thereby pro-
viding theoretical basis for reasonable
irrigation of P. volubilis plants in the
field during the dry season.
Materials and Methods
Materials
The test was carried out in the
greenhouse of Xishuangbanna Tropi-
cal Botanical Garden (21°56′ N, 101°
1′ E; altitude 560 m) during November
20, 2013 to January 11, 2014. There
were fly nets around the greenhouse,
and the average temperature inside
the greenhouse was about 22 ℃ . The
mature seeds of P. volubilis were
nursed in sandy soil. The growth-uni-
form young seedlings of P. volubilis, in
height of about 20 cm, were trans-
planted to pots (inner diameter 26 cm,
height 23 cm) with 7 kg soil. The basic
physical and chemical properties of
the tested soil were as follows: organic
matter 18.39 g/kg, available nitrogen
110 mg/kg, available phosphorus5.98
mg/kg, available potassium 98 mg/kg.
Methods
Experimental design
A total of three factors were de-
signed, i.e., irrigation mode, irrigation
amount and fertilization. The irrigation
modes included conventional irrigation
and alternative partial root-zone irriga-
tion (RD). The irrigation amount cov-
ered three levels, i.e., normal supply
(NS, 100% of transpiration amount;
transpiration amount referred to the
weight difference of every pot every
two days), mild drought (MD, 75% of
transpiration amount) and severe
drought (SD, 50% of transpiration
amount).For fertilization, nitrogen fertil-
ization (N, along with urea, 5 g/plant)
and no fertilization treatments were
used. A total of seven treatments
(Table 1) were designed with 20 pots
for every treatment. In the RD treat-
ment group, the pot was equally divid-
ed into two parts with a plastic plate,
and the roots of every P. volubilis plant
were equally divided into the two parts;
the roots in the two parts were irrigated
once alternatively every two weeks.
After 10-d adaptation with full irriga-
tion, the measurement was started.
The pots in all the treatment groups
were covered with plastic film to re-
duce soil evaporation. The water sup-
ply was strictly controlled, and the irri-
gation was carried out by hand. The
pots were weighed and irrigated once
every two days. The irrigation amount
of every pot was recorded every time.
Total irrigation amount (I) of every pot
referred to the sum of irrigation
amounts during the test. The nitrogen
fertilizer was dissolved in irrigation wa-
ter and applied diagonally uniformly
around the seedlings. Throughout the
experimental period, the nitrogen fer-
tilizer was applied four times.
Measurement
When the test began, total five
young seedlings were selected ran-
domly for every treatment. They were
dried at 105 ℃ for 30 min, and then
at 70 ℃ to constant weight (W1). Be-
fore the test ended, the net photosyn-
thetic rate (Pn), transpiration rate (Tr),
stomatal conductance (Gs) and inter-
cellular CO2 concentration (Ci), of fully
expanded mature leaves of P. volu-
bilis were measured withLi-6400
portable photosynthesis system dur-
ing 8:30 -11:00 of a sunny day. The
instantaneous water use efficiency
(WUEi, Pn/Tr) was calculated. When
the test ended, total five plants were
sampled randomly for every treatment.
The fresh blades were scanned with-
CanoScan4400F scanner, and their
leaf area indexes were calculated us-
ing ImagJ software. The five plants in
every treatment were divided into
roots, stems and leaves, which were
dried at 105 ℃ for 30 min and then
at 70 ℃ to constant weights (W2, W3
and W4). The nitrogen content (N, %)
in dried leaves was measured with
TOC/TN analyzer.
Root cap ratio (g/g) = W2/(W3 +
W4);
Photosynthetic N utilization effi-
ciency (μmol/(mol2·s·N) =Pn/N;
Irrigation water use efficiency
(WUE, g/L) = (W2+W3+W4-W1)/I.
Data statistics and analysis
The data were analyzed using
SPSS 13.0. ANOVA tests were con-
ducted for the no fertilization treat-
ments, and the multiple comparisons
were conducted with least significant
difference (LSD) method (α =0.05).
Tow-way ANOVA test was conducted
for the severe drought treatment. All
figures were drawn using SigmaPlot.
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Agricultural Science & Technology 2016
Different lowercase letters indicate significant differences at the 0.05 level.
Fig.1 Comparisons of biomass, root cap ratio and leaf area index of P. volubilis seedlings among different treatment groups
Results and Analysis
Growth and biomass
Biomass, as one of the most im-
portant indicators, could intuitively re-
flect the growth of plants. As shown in
Fig.1-A, no significant differences were
found in biomass of young P. volubilis
seedlings among the no fertilization
treatment groups (P>0.05). With the
decreased irrigation amount, the
biomass of young seedlings was re-
duced. The biomass of young
seedlings in the NS treatment group
was highest, and it was 0.31 g (27%)
higher than that all in RD treatments.
Water shortage affected the biomass
accumulation in young P. volubilis
seedlings. When the irrigation amount
was equivalent, the biomass of young
seedlings under alternative partial
root-zone irrigation was higher than
that under conventional irrigation.
Compared with that in the MD treat-
ment, the biomass in the MD +RD
treatment was higher by 15%, and that
in the SD+RD was higher by 18%. Be-
sides, there was no significant differ-
ence among NS, MD+RD and SD+RD
treatments, and the seedling biomass
in the NS treatment was higher than
those in the MD +RD and SD +RD
treatments by 5% in average. In the
case of limited irrigation water, alter-
native partial root-zone irrigation could
alleviate water stress to some extent,
and ensure biomass accumulation.
Under severe drought, there was no
significant difference in seedling
biomass between irrigation mode and
fertilization (P>0.05). It suggested that
under severe drought condition, irri-
gation mode and N fertilization showed
little effect on biomass accumulation in
P. volubilis seedlings. The seedlings
biomass in the SD +RD +N treatment
group was 7% higher than that under
severe drought, but was 12% lower
than that in the SD +RD treatment
group. It indicated that under the con-
dition of severe drought, N fertilization
affected the biomass accumulation in
young P. volubilis seedlings.
Root is the main vegetative organ
of plants to absorb water and nutrients.
Root cap ratio reflects the distribution
of biomass between ground and un-
derground parts of plants. The vari-
ance analysis (Fig.1-B) showed that
there were no significant differences in
root cap ratio among the no fertilization
treatment groups (P >0.05).The root
cap ratio under severe drought was a
little higher. It indicated that under the
condition of severe drought, biomass
was preferentially allocated to the
roots of plants. Under severe drought,
significant difference was found in root
cap ratio between irrigation mode and
fertilization (P<0.05), but their interac-
tion was insignificant (P>0.05). In aver-
age, the root cap ratio under N fertil-
ization was about 30% higher than that
when no fertilizer was applied. Under
serious water shortage, alternative
partial root-zone irrigation and N fertil-
ization significantly promoted the
transport of nutrients to seedling roots,
increased root cap ratio and root
biomass, and increased root ability to
absorb nutrients and water, thus im-
proving drought resistance.
Leaves are the main photosyn-
thetic organ, and leaf area index re-
flects the growth conditions of leaves
of young seedlings. The variance anal-
ysis (Fig.1-C) showed that water sup-
ply significantly affected seedling leaf
area index (P<0.05). The leaf area in-
dex in the NS treatment was 25% and
42% higher than those in the MD and
RD treatments respectively, but was
only 7% and 27% higher than those in
the MD +RD and SD +RD treatment
groups. It indicated that drought stress
inhibited leaf growth, but alternative
partial root-zone irrigation alleviated
the adverse effects of drought stress
on the growth of seedling leaves.
There was significant difference in leaf
area index between irrigation mode
and fertilization (P<0.05), but their in-
teraction was insignificant (P >0.05).
The leaf area index in the SD +RD
treatment group was 38% higher than
that in the SD+RD+N treatment group.
Under the condition of alternative par-
tial root-zone irrigation, fertilization was
not conducive to the leaf growth of P.
volubilis under severe water stress.
Photosynthetic physiology
As shown in Fig.2, there was no
significant difference in photosynthetic
rate (P>0.05), but there were signifi-
cant differences in transpiration rate
and stomatal conductance (P <0.05)
between normal water supply and mild
drought treatments.The transpiration
rate and stomatal conductance in the
MD +RD treatment group were 33%
and 36% lower than those under nor-
mal water supply, and were 9% and
18% lower than those under mild
drought. It indicated that with de-
creased irrigation amount, transpira-
tion rate and stomatal conductance
were reduced as well. However, under
the condition of mild drought, alterna-
tive partial root-zone irrigation signifi-
cantly reduced stomatal conductance
and transpiration rate, instead of pho-
tosynthetic rate. The photosynthetic in-
dexes in the SD+RD treatment group
were significantly lower than in the oth-
er irrigation treatments. The variance
analysis showed that irrigation mode
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Agricultural Science & Technology2016
Different lowercase letters indicate significant differences at the 0.05 level.
Fig.2 Comparisons of photosynthetic physiology and N use efficiency of young P. volubilis
seedlings among different treatment groups
Different lowercase letters indicate significant differences at the 0.05 level.
Fig.3 Comparison of water use efficiency in young P. volubilis seedlings among different
treatment groups
and fertilization showed no significant
effects on photosynthesis of P. volu-
bilis (P>0.05). It suggested that water
stress significantly inhibited the photo-
synthesis process, and its effect on P.
volubilis seedlings was greater than
thoseof irrigationmodeand fertilization.
There were no significant differ-
ences in leaf N content among the no
fertilizer treatment groups (P >0.05),
but the average leaf N content in the
fertilization treatment groups was 10%
in average higher than that in the no
fertilizer treatment groups. The ab-
sorption of water and N fertilizer was
independent for plants, and drought
stress did not affect the absorption and
utilization of N fertilizer in P. volubilis
seedlings. Instantaneous nitrogen use
efficiency reflects (PNUE) the status of
N physiological utilization in crop
blades, and it is the main factor deter-
mining plant growth and leaf nitrogen
productivity. The variance analysis
showed that the PNUE under severe
drought was significantly lower than
those in the other treatments (P<0.05),
indicating water shortage affecting
PNUE. Under severe drought condi-
tion, fertilization had a significant effect
on N photosynthetic rate (P <0.05),
however, the interaction between irri-
gation mode and fertilization was in-
significant (P >0.05). The average
PNUE of the fertilization treatment
groups was 22% lower than that of no
fertilizer treatment groups. The PNUE
was negatively related to leaf nitrogen
content.
Leaf instantaneous water use effi-
ciency (WUEi) only reflects the status
of water utilization in leaves within a
certain time, and the relationship be-
tween the results of instantaneous
and long-term overall measurement is
unclear[14]. Therefore, WUEi is not suit-
able for reflecting water use efficiency
throughout the test period.
As shown in Fig.3, there were no
significant differences in WUEi among
the irrigation treatments except MD +
RD and NS treatment [the WUEi in the
MD +RD treatment group was about
25% higher than that in the NS treat-
ment group]. For MD+RD treatment, it
reduced the transpiration efficiency
without reducing photosynthetic rate.
This might be because that drought
stress force P. volubilis close the
stomas, and its luxury transpiration
was reduced, thus the WUEi was im-
proved. In the SD treatment, the inter-
actions between irrigation mode and
fertilization on WUEi was insignificant
(P >0.05). However, the WUEi in the
fertilization treatment groups was
higher, which might be due to fertiliza-
tions improving leaf nitrogen content
and photosynthetic rate.
Irrigation water use efficiency
Fig.3 showed that there were sig-
nificant differences in water use effi-
ciency (WUE) of young P. volubilis
seedlings among different irrigation
treatments (P <0.05). With the de-
crease of irrigation amount, the WUE
of young seedlings increased signifi-
893
Agricultural Science & Technology 2016
cantly. The WUE in the SD+RD treat-
ment group was highest, and it was
about 60% higher than that under
normal water supply. Maybe P. volu-
bilis seedlings had small demand for
water or alternative partial root-zone
irrigation contributed to improving the
WUE, especially under severe drought
condition. Irrigation mode had signifi-
cant effect on WUE in P. volubilis
seedlings (P<0.05), but the interaction
between irrigation mode and fertiliza-
tion was insignificant (P >0.05). The
WUE in the SD +N treatment group
was higher than that in the SD+RD+N
treatment group. It indicated that under
severe drought stress condition, the
mode of alternative partial root -zone
irrigation could reduce transpiration
without affecting photosynthesis,
thereby maximizing the WUE. Howev-
er, the factor of N fertilization failed to
show the affect.
Conclusions and Discus-
sion
The results of this study showed
thatalternative partial root-zone irriga-
tion could improve water use efficien-
cy and alleviate drought stress to
some extent by adjusting the leaf
stomas, and it is an effective water-
saving irrigation technology. Com-
pared with those under full irrigation,
the biomass and water consumption
under alternative partial root-zone ir-
rigation were reduced by 5% and
75%, respectively, and the water use
efficiency was increased by 60%. This
conclusion was consistent with the
findings in rice, potatoes, corn and
other field crops, and grapes, peach
and other fruit trees [13,15-18]. Alternative
partial root-zone irrigation can improve
water use efficiency. This might be
due to the stress signals generated by
the root under drought stress, such as
ABA. When the signals are transferred
to the aboveground parts, the opening
or closure of leaf stomas is regulated.
Stomatal conductance shows linear
relationships with transpiration water
consumption, but shows non-linear
relationship with photosynthetic rate.
When the stomatal conductance de-
clines gradually from the maximum,
luxury transpiration is reduced signifi-
cantly, but the photosynthesis is affect-
ed lightly. This difference provides a
theoretical basis for improving water
use efficiency of plants through regu-
lating stomatal activities [9,19]. In this
study, compared with those under full
irrigation, the stomatal conductance,
photosynthetic rate and transpiration
rate under alternative partial root-zone
irrigation were reduced by 29%, 20%
and 38%, respectively, but the instan-
taneous water use efficiency was in-
creased by 19%. In average, the water
use efficiency in the extreme drought
treatment groups was higher than that
in the other treatment groups. The wa-
ter use efficiency in the SD+RD treat-
ment group was highest, and it was
four times higher than that under nor-
mal water supply. This was consistent
with the result in citrus[20].
Studies suggest that under
drought conditions, ABA, produced by
roots, is transported to shoots, and it
can regulate the closure of leaf stoma-
ta, reduce the expansion rate of blades,
reduce leaf growth, and allocate more
assimilates to the roots, thereby pro-
moting root growth, but inhibiting shoot
growth; thus, the root cap ratio is regu-
lated[10]. In this study, alternative partial
root-zone irrigation did not significantly
increase the root cap ratio, but the root
cap ratio in the SD +RD treatment
group was significantly higher (about
30%) than those in the other treatment
groups. This conclusion was inconsis-
tent with the findings of some previous
studies[21-22], which probably be caused
by the differences in experimental
conditions, plant species and growth
state. In addition, alternative partial
root-zone irrigation could alleviate the
drought to some extent. However, un-
der the condition of long-term severe
drought, alternative partial root-zone
irrigation allocated more biomass to
the roots, greatly reduced the leaf area
index, and decreased photosynthetic
indicators. Thus, the growth and de-
velopment of P. volubilis seedlings
was inevitably affected.
Under the condition of limited wa-
ter, appropriate application of nitrogen
can improve photosynthetic charac-
teristics of plants, promote plant
growth, and improve water use effi-
ciency and drought tolerance of plants
to some extent[6, 23]. In the SD+N treat-
ment group, the photosynthetic rate
was increased by 6% , the biomass
was increased by 11%, and the water
use efficiency was increased by 25%.
However, the biomass in the SD+RD+
N treatment group was reduced by
12% compared with that in the SD+N
treatment group. Under the conditions
of severe drought and alternative par-
tial root-zone irrigation, nitrogen appli-
cation adjusted the distribution of
biomass and concentrated most of the
biomass to roots of P. volubilis
seedlings. Thus, the root cap ratio and
root biomass were all increased by
30% in average, so as to enhance the
drought tolerance of young P. volubilis
seedlings. However, the leaf area in-
dex was reduced by 38% , the photo-
synthetic rate was reduced by 9%, and
the total biomass was reduced by
18%. Alternative partial root-zone irri-
gation was not suitable to be matched
with nitrogen fertilization under severe
drought conditions. Photosynthetic ni-
trogen use efficiency (PNUE), as an in-
dex for evaluating the relationship be-
tween Pn and leaf nitrogen content, is
commonly used in studies on vegeta-
tion adaptation and plant population
evolution. In crops, PNUE is found
closely related to photosynthetic char-
acteristics, yield and nitrogen use effi-
ciency[24-26]. In the SD+N and SD+RD+
N treatment groups, the PUNE was
negatively related to leaf nitrogen
content, which was consistent with
previous studies[27-28]. This might be be-
cause that the nitrogen was assigned
to the leaf photosynthetic apparatusto
constitute cell walls. As this experi-
ment was conducted under the condi-
tion of pot cultivation in the green-
house, the effect of implementation of
alternative partial root-zone irrigation in
the field conditions needs to be ex-
plored. In the field, the effects of het-
erogeneous rainfall and the duration of
drought stress should also be taken
into account when water-saving irriga-
tion technologies are adopted.
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