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Growth and yield responses of Sacha Inchi (Plukenetia volubilis) plants to

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Journal of Horticultural Science & Biotechnology (2014) 89 (2) 201–207

Planting density and fertilisation independently affect seed and oil

yields in Plukenetia volubilis L. plants

By C. YANG1,2, D. Y. JIAO1,Y. J. GENG1,2 C. T. CAI1 and Z. Q. CAI1*

1
Key Laboratory of Tropical Plant Resource Science, Xishuangbanna Tropical Botanical Garden,
Chinese Academy of Sciences, Mengla, Yunnan 666303, P. R. China
2
Graduate University of the Chinese Academy of Sciences, Beijing 100049, P. R. China
(e-mail: zhiquan.cai@126.com) (Accepted 2 December 2013)

SUMMARY
To investigate the effects of planting density and fertiliser level on some important agronomic traits of Plukenetia
volubilis, a perennial oilseed species, a field experiment with four replications in a completely randomised block design
was conducted in a tropical region of China in the 2012-2013 growing season. Planting density (1,666, 2,500, 4,444,
5,000, or 10,000 plants ha–1) was assigned to the main plots and a 1:1:1 (w/w/w) NPK fertiliser at 0, 50 100, 150, or 200
kg ha–1 was assigned factorially to the sub-plots. The results showed that neither planting density nor the rate of
fertilisation affected seed size (dry matter) and the phenological development of P. volubilis plants, including the times
of initial flowering and maturity, or the pattern of fruit ripening. Planting density, rather than fertiliser level,
significantly affected seed oil content, but a non-linear effect was found. Across the different treatments and sampling
times, seed size and oil content had relatively high and constant heritabilities indicated by low coefficients of variation.
Total seed yields and total oil yields per plot over the growing season ranged from 1,340 - 2,486 kg ha–1, and from 501
- 899 kg ha–1, respectively, with a quadratic response to planting density at all fertiliser levels. Total seed and oil yields
increased continuously with the increasing levels of fertiliser used. The absence of any planting density ! fertiliser level
interaction on total seed and oil yields suggested that planting density and fertilisation had independent effects, and
that seed or oil yield responses to planting density were not significantly affected by fertiliser level. We concluded that
oil production in P. volubilis plants required high levels of fertiliser and, regardless of fertiliser level, a planting density
of approx. 4,444 plants ha–1 was required to ensure maximum yield in the field.

P lanting density has important effects on the

vegetative and reproductive development of crops
(Ciampitti and Vyn, 2011; Dong et al., 2012). A high
premature senescence, and decreased fruiting, thus
limiting the yield potential (Zhang et al., 2012).
Although it has been widely reported that the
planting density is commonly used to increase crop application of fertiliser (especially N) increased plant
yields. However, if the planting density is too high, this growth and crop yields in the field (Jackson, 2000;
reduces the availability of resources (i.e., incident Rotundo and Westgate, 2009), an over-dose of fertiliser
radiation, water, and nutrients) per plant in the growing can promote excessive vegetative development at the
season and causes a decline in yield per plant which may expense of fruit yield, and probably also delays maturity
not be offset by the increase in planting density (Cheema et al., 2001). The independent effects of
(Andrade et al., 1999). A high planting density in maize planting density or level of fertilisation on crop yield
increased total dry matter production and decreased the and yield components have been well-documented
harvest index (HI). The optimum planting density (Zhang et al., 2012; Cai et al., 2013), but their interactions
reflected a balance between these two effects have been relatively less well-studied, especially in the
(Tollenaar, 1989). Although the effects of planting field.
density on crop yield and on various yield components Plukenetia volubilis L. is native to South America and
have been studied in many horticultural crops, the is a promising new oilseed crop in the family
results have varied greatly between species. In many Euphorbiaceae. P. volubilis is a perennial woody vine
species, increased planting densities, up to an optimum that produces seeds with a high oil content [40 – 60%
level, resulted in increased yields (Brahim et al., 1998). (w/w)] which exceeds the quality characteristics of
In some species, a lower planting density prolonged the existing oils used for human consumption worldwide.
seed-filling period, and thus increased seed size (Rogers P. volubilis seed oil is one of the richest sources of
and Lomman, 1988). The seed yield per unit area, in unsaturated fatty acids and its domestic, industrial,
response to planting density, depends not only on the medicinal, and cosmetic industry uses have increased the
species and environmental conditions, but also on the global value of this crop (Cai et al., 2011). P. volubilis
level of fertiliser applied and other agronomic practices plants do not exhibit winter dormancy. They grow
(Ciampitti and Vyn, 2011). Limited amounts of available continuously in tropical regions and therefore flower and
soil nutrients lead to weak plant development, fruit almost continuously throughout the year. Each fruit
is a capsule (ca. 4 – 7 cm in diameter), consisting of four-
*Author for correspondence. to-seven pods, with one seed per pod.
202 Plukenetia volubilis responses to planting density and fertiliser

The processes determining the quantity and quality of Measurements

oil in P. volubilis seed are variable and depend on several Twenty small fruit (diameter < 0.5 cm) in each sub-
environmental conditions and agronomic factors (Cai, plot were tagged at random on 19 November and on 12
2011; Jiao et al., 2012). In addition, P. volubilis plants have December 2012 in only the highest (200 kg ha–1) and
the capacity to develop new reproductive structures in lowest (0 kg ha–1) fertiliser level sub-plots at each
response to an increase in available resources (Cai, 2011; planting density. Approx. 1.5 months later, the numbers
Jiao et al., 2012; Cai et al., 2012; 2013). Therefore, we of mature fruit were counted and the “fruit abortion
predict that there is potential to increase seed yields in percentage” was calculated as the proportion of empty
P. volubilis plants by suitable agronomic management fruit compared to the total number of fruit tagged.
practices. The stem diameter of each plant was measured 2 cm
In a preliminary study, we found that a relatively high above soil level, using calipers, to measure plant growth
planting density was required to optimise seed yields in at the end of April 2013. Five-to-six plants were selected
this new oilseed crop (Cai et al., 2013). To date, no at random in each sub-plot, and the dates of first bloom
optimum planting density or rate of fertiliser application and mature fruit set were recorded for each plant.
has been established to achieve the maximum levels of Mature fruit from all P. volubilis plants were harvested
seed or oil production. Therefore, in the present study, a seven-times, by hand, in each replicate sub-plot,
field experiment was conducted to investigate the throughout the 5-month period of fruit ripening. The
combined effects of planting density and rate of fertiliser total dry mass (DM) of fruit per plot was measured at
application on the reproductive traits, total seed and oil each harvest. Also at each harvest, sub-samples of
yields, yield components, and seed quality during a single mature fruit were peeled and the DM (size) of all seeds
growing season at Xishuangbanna, a tropical region of per plot was recorded.
China. The overall goal was to provide a better Seed oil contents were determined using a Minispec
understanding of planting density and fertiliser mq-one Seed Analyser (Bruker Optik GmbH, Ettlingen,
management for this species at both the local and Germany). A calibration curve was obtained using
regional levels, and thus to increase seed and oil yields known reference samples of the oil extracted from
for commercial-scale oil production. P. volubilis seeds. As the average DM of seeds accounted
for 53.4% of the total fruit DM, and did not differ across
all sampling dates and treatments (n = 150), oil yield at
MATERIALS AND METHODS each harvest was estimated as follows:
Experimental site, plant material, and experimental
treatments Oil yield (kg ha–1) = Mature fruit DM (kg ha–1) ! 0.534 !
Seeds of P. volubilis were sown in a nursery in Seed oil content.
February 2012. When the seedlings were approx. 20 cm The total oil yield (ha–1) throughout the growing
tall, in late April 2012, uniformly-sized seedlings were season was then calculated by adding the values from
selected and cultivated at four open sites at the each harvest.
Xishuangbanna Tropical Botanical Garden of the
Chinese Academy of Sciences (21°56’N, 101°15’E; 560 m Statistical analysis
asl). The soil was a red–brown type and soil samples were All data are presented as mean values ± standard
collected in October 2011. The characteristics of the top deviations (SD). Seed size (DM), oil content, and seed
(0 – 20 cm) layer of soil were: pH 5.42; organic carbon and oil yields between the different sampling times were
5.65% (w/v); total nitrogen 0.34 g kg–1; available N 46.0 analysed by one-way ANOVA and means were compared
mg kg–1; available P 14.1 mg kg–1; and available K 22.0 mg by Tukey test at P ≤ 0.05. We used two-way ANOVA to
kg–1. The climate at Xishuangbanna is dominated by the compare fruit set percentages, seed traits, and total seed
Southwest monsoon with two distinct seasons (a wet and oil yields between the different planting densities,
season from May to October, and a dry season from fertiliser levels, and their interactions across the different
November to April; Cao et al. 2006). The average annual sampling times. We then used LSD contrasts to examine
temperature is 22.9ºC and the mean annual precipitation whether each trait differed between planting densities
is 1,500 mm, of which approx. 85% occurs in the wet within or between fertiliser levels. The data were tested
season. for normality and homogeneity of variance and, when
The field experiment was conducted using a necessary, were log10-transformed before analysis. The
randomised complete block design with four replicates variability of each trait for each factor (planting density
of each treatment. Planting density was assigned to the or fertiliser level) was quantified using coefficients of
main plots with five different intra- and inter-row variation [CV; i.e., the standard deviation (SD) divided by
spacings: 3.0 m ! 2.0 m; 2.0 m ! 2.0 m; 1.5 m ! 1.5 m; the mean of the characteristic]. All statistical analyses
2.0 m ! 1.0 m; or 1.0 m ! 1.0 m, resulting in planting were conducted using SPSS software Version 13.0 (SPSS
densities of 1,666, 2,500, 4,444, 5,000, or 10,000 plants Inc., Chicago, IL, USA).
ha–1, respectively. Fertilisation rates were assigned to the
sub-plots and consisted of five levels (0, 50, 100, 150, or
200 kg ha–1) of a 1:1:1 (w/w/w) mix of NPK spread in an RESULTS AND DISCUSSION
approx. 1.0 m-wide zone in June 2012 (the wet season). Plant reproductive pattern
In total, there were 100 sub-plots, each approx. 40 m in P. volubilis plants in all treatments started flowering
length. between 18 – 20 June 2013. Neither planting density nor
Since P. volubilis is a liana species, all plants were rate of fertilisation affected the time of initial flowering
supported to a height of 1.6 m using steel wires. or the first harvest date, which was contrary to previous
 C. YANG, D. Y. JIAO, Y. J. GENG, C. T. CAI and Z. Q. CAI 203

TABLE I
Summary of two-way ANOVA to evaluate the effects of planting density,
rate of fertilisation, and their interaction on seed and oil traits and mean
values of each trait under different treatments
Seed oil
Seed yield Seed oil yield content Seed size
Factor (kg ha–1) (kg ha–1) [%(w/w)] (g seed–1)
Two-way ANOVA (significance level)§
PD‡ *** *** *** ns
F ** * ns ns
PD ! F ns ns ns ns
Treatment means
PD (plants ha–1)
10,000 1,799.0† bc 666.2 b 37.4 a 1.32 a
5,000 1,986.6 ab 725.9 ab 36.6 b 1.35 a
4,444 2,154.6 a 786.1 a 36.5 b 1.36 a
2,500 1,549.2 d 572.5 c 37.1 a 1.37 a
1,666 1,584.8 cd 564.7 c 35.6 c 1.38 a
‡
CV (%) 14.3 14.5 1.9 1.70
Fertilisation (kg ha–1)
0 1,626.6†c 598.8 c 36.7 a 1.35 a
50 1,745.8 c 640.5 c 36.7 a 1.34 a
100 1,812.7 ab 663.6 ab 36.5 a 1.36 a
150 1,860.4 ab 673.1 ab 36.6 a 1.35 a
200 2,028.6 a 739.4 a 36.6 a 1.36 a
CV (%) 8.2 7.8 0.23 0.62
†
Values are means (n = 4). Values followed by different lower-case
letters in each column indicate significant difference at P ≤ 0.05 by the
LSD test.
‡
PD, planting density; F, fertilisation; CV, coefficient of variation.
§
ns, not significant (P > 0.05); or significant at *, P ≤ 0.05; **, P ≤ 0.01;
***, P ≤ 0.001.

affected fruit set in P. volubilis plants only at the early

reproductive stage (November) with the highest fruit set
values observed at medium planting densities
(Figure 1 A). At higher plant densities, increased
crowding could reduce plant growth around the critical
seed-filling period, resulting in fruit abortion (Andrade
et al., 1999), and thus reduced kernel numbers and seed
yields per plant. There are reports that plant density
affects fruit set in a large number of wild species in a
wide variety of habitats (Stephenson, 1981; Zorn-Arnold
and Howe, 2007). Fertiliser application generally
increased the fruit set in P. volubilis plants at both the
FIG. 1
Fruit set percentages in P. volubilis plants cultivated at five different early and late reproductive stages (Figure 1 A,B), which
planting densities and two rates of fertilisation (LF or HF) in November agreed with the findings of our previous study (Jiao et al.,
(Panel A) and December (Panel B) 2012. PD, planting density; F,
fertiliser. LF, low fertilisation rate (0 kg ha–1); HF, high fertilisation rate
2012). A high fruit set percentage in P. volubilis plants
(200 kg ha–1). Significant at *, P ≤ 0.05; **, P ≤ 0.01; ns, not significant. under high fertiliser levels required an increased
photosynthetic input and thus reduced carbohydrate
availability (Stephenson, 1981).
results that long-term exposure to shade delayed the Over the 5-month period of production, the patterns
initial date of flowering in P. volubilis plants (Cai, 2011). of both mature seed and oil yields at the sub-plot level
The same initial flowering times with increases in plant were similar between the different treatments, with the
density and rate of fertilisation would not necessarily highest values observed in the middle of March
lead to a delay in the time of harvest. Neither planting (Figure 2 C,D). Seeds that matured later (i.e., after
density nor rate of fertilisation affected the phenological March) had a relatively large size, whereas their seed oil
development of P. volubilis plants. This was due to the content was not affected by sampling date
fact that the time from sowing to flowering (or seed (Figure 2 A,B).
maturity) within one season in plants with no
requirement for vernalisation is largely determined by Seed oil content and seed size (DM)
their responses to temperature and photoperiod The influence of planting density on the oil content of
(Olesena et al., 2012). oilseed crops has been controversial. For example, there
As a wind-dispersed species with well-developed have been reports of an increase (Zhang et al., 2012), a
reproductive organs (i.e., flower and fruit numbers; Jiao decrease (Momoh and Zhou, 2001), or no effect (Leach
et al., 2012), seed production in P. volubilis depends et al., 1999) on the oil contents of winter oilseed rape
mainly on the availability of resources to make seeds seed (Brassica napus L.) with increasing planting density.
(Zorn-Arnold and Howe, 2007). There were no planting There have also been reports that the oil contents of
density ! fertiliser interactions on fruit set in P. volubilis castor bean, (Ricinus communis; Soratto et al., 2012) and
plants at the early (November) or late (December) sunflower (Helianthus annuus; Narwal and Malik, 1985)
reproductive stages (Figure 1 A,B). Planting density seeds were not affected by planting density. In the
204 Plukenetia volubilis responses to planting density and fertiliser

present study, planting density had a significant, but non-

linear affect on the oil content of P. volubilis seeds
(Table I). Across the different treatments and sampling
times, seed oil contents ranged from 33.9 – 37.4% (w/w),
which was comparable to previous reports (Cai et al.,
2012; Jiao et al., 2012).
It is likely that increased nutrient stress during the
seed-filling period resulted in seeds with a reduced oil
content (Rotundo and Westgate, 2009). However, seed
oil contents decreased with increasing levels of
fertilisation in many studies (Jackson, 2000; Cheema
et al., 2001; Rathke et al., 2005). This might be due to the
reduced availability of carbohydrates for the synthesis of
oils under high N supply. However, oil contents did not
necessarily correlate inversely with protein contents
across different environments (Cai et al., 2012) because
the biosynthesis of storage proteins and oils in
developing oil seeds are two independent processes, and
the genes that control oil and protein synthesis are
expressed differently (Votlker and Kinney, 2001). The
rates of fertilisation applied here did not affect the oil
contents of P. volubilis seed across all sampling dates
(Table I), which agreed with results from canola and
soybean (Hocking et al. 1997; Rotundo and Westgate,
2009).
A lower planting density would result in a prolonged
seeding-filling period, thus increasing seed size (Leach
et al., 1999). However, neither planting density nor rate
of fertilisation affected seed size (i.e., seed DM) in
P. volubilis plants across the different sampling dates
(Figure 1 A), confirming results in canola (Angadi et al.,
2003). In addition, similar-sized seeds across the different
treatments revealed that an increased number of seed
(fruit) ha–1, rather than individual seed size, was mainly
responsible for the total seed yield throughout the
growing season (Cai et al., 2013). The coefficients of
variation in oil content and seed size in response to
planting density and rate of fertilisation were small (0.23
– 1.90%), indicating that these traits had a relatively high
and constant heritability in P. volubilis plants (Soratto
et al., 2012). Moreover, the low variation in seed oil
content indicated that seed quality did not vary greatly
across the different treatments throughout the 5-month
fruit-ripening period.

Total seed and oil yields

There were no planting density ! fertiliser interactions
on total seed or oil yields. Thus, planting density and rate
of fertilisation were assumed to have independent
effects. P. volubilis plants showed a similar slight increase
in total seed or total oil yields in response to fertiliser
level with increased planting density, while the response
to planting density was not greatly affected by fertiliser
level. This contradicted observations in maize (Asim
et al., 2013) and corn (Dong et al., 2012), in which the
interaction of plant density and nitrogen level had
significant effects on crop yield. Both planting density
and fertiliser level significantly affected total seed and oil
FIG. 2 yields (Table I; Figure 2; Figure 3). Moreover, the
Mature seed size (Panel A), seed oil content (Panel B), seed yield coefficients of variation were higher in response to
(Panel C), and seed oil yield (Panel D) in P. volubilis plants during one
growing season cultivated at different levels of fertilisation combined planting density than to fertiliser level, indicating that
with a planting density of 4,444 plants ha–1 as an example. Mean values planting density may be more important as a limiting
(n = 4) with different lower-case letters in each Panel indicate significant
differences between fertilisation levels for this planting density at factor than fertiliser level during the production of seeds
P ≤ 0.05. ns, not significant. and oil in P. volubilis plants (Diepenbrock, 2000).
 C. YANG, D. Y. JIAO, Y. J. GENG, C. T. CAI and Z. Q. CAI 205

FIG. 3
Total seed yield (Panel A), total seed oil yield (Panel B), seed oil content
(Panel C), and mature seed size (Panel D) in P. volubilis plants FIG. 4
cultivated at five different planting densities and five rates of Relationships between planting density and total oil yield (Panel A) or
fertilisation pooled across the different sampling times. Mean values (n total seed yield (Panel B) at five different rates of fertilisation
= 4) with a different lower-case letter in each Panel indicate significant throughout a growing season, and between stem diameter and total seed
differences between fertilisation levels within each planting density at yield (Panel C) in P. volubilis plants across five different fertilisation
P ≤ 0.05. ns, not significant. treatments.

Total seed and oil yields over the growing season at in planting density to 4,444 plant ha–1 was an effective
the sub-plot level ranged from 1,340 – 2,486 kg ha–1, and means to increase seed and oil yields in P. volubilis
from 501 – 899 kg ha–1, respectively, among the different plants, after which point both total seed and oil yields
treatments. The increase in total seed yield was declined significantly (Table I; Figure 4 A,B). Regression
accompanied by an increase in plant growth (i.e., stem analysis indicated that the relationships between
diameter; Figure 4 C), indicating that the resources planting density and total seed or oil yields at any
demanded by the seed are controlled mainly by the fertilisation level were best expressed by quadratic
amount of stored resources (i.e., carbohydrates) in the equations (R2 = 0.76 – 0.89; all P ≤ 0.01; Figure 4 A,B),
maternal plant during the reproductive stage. The namely:
highest yields of both seed and oil occurred at the Y = a + b D + c D2
medium planting density combined with the highest rate
of fertilisation (Table I; Figure 3; Figure 4). An increase where Y was the total seed yield or the total oil yield (kg
206 Plukenetia volubilis responses to planting density and fertiliser

ha–1), D was the planting density (plants ha–1), and a total seed and oil yields increased continuously with
(intercept), b (linear coefficient), and c (quadratic increasing levels of fertilisation (Table I; Figure 2),
coefficient) were parameters. suggesting that P. volubilis plants require relatively high
Based on these quadratic equations, the estimated fertiliser levels to achieve maximum yields in the field. A
optimum planting density for the maximum total seed or strong positive correlation was found between total seed
oil yields did not vary greatly at all fertiliser levels. and oil yields across all sampling dates (r = 0.99; P ≤
Therefore, a planting density of approx. 4,444 plants ha–1 0.001). Thus, oil production in P. volubilis plants is
is required to ensure the optimum yield of P. volubilis determined primarily by the maximum attainable seed
plants, regardless of fertiliser level. Seed and oil yields yield, but not by seed oil content (Sidlauskas and
commonly increased curvi-linearly with an increase in Bernotas, 2003; Rathke et al., 2005).
planting density in other oilseed crops such as winter In conclusion, neither planting density nor rate of
oilseed rape (Leach et al., 1999; Momoh and Zhou, 2001; fertilisation affected the phenological development of
Sidlauskas and Bernotas, 2003), Zea mays L. (Ciampitti P. volubilis plants. Planting density and fertilisation
and Vyn, 2011), and in food crops such as faba bean affected seed and oil yields in P. volubilis plants
(Vicia faba L.; López-Bellido et al., 2005), and in cotton independently. A planting density of approx. 4,444 plants
(Dong et al., 2012). ha–1 was required to ensure maximum seed and oil yields,
As the partitioning of DM to reproductive sinks regardless of fertiliser level. P. volubilis plants require
during flowering, and the DM: total DM ratio of seeds relatively high rates of fertilisation application to achieve
were reduced at harvest by shortages of N (Ciampitti and their maximum seed and oil yields. Future research
Vyn, 2011), positive impacts of the application of should focus on a detailed analysis of the optimum rate
fertiliser on seed yields in oilseed crops such as winter of application of fertiliser to achieve the highest yield of
oilseed rape and soybean have been reported (Jackson, P. volubilis plants and to mitigate losses of nutrients in
2000; Rathke et al., 2005). In contrast, some authors have the field.
reported a plateau or even a reduction in seed yields at
high rates of N-fertiliser (Hocking et al., 1997; Cheema This work was supported financially by the Chinese
et al., 2001). The optimum seed yield in oilseed crops Academy of Sciences (Grant Nos. KSCX2EWQ17 and
under a particular fertiliser level depends on KSCX2EWZ15) and by the National Science Foundation
environmental conditions (Jackson, 2000). In our study, in China (Grant Nos. 31170641 and 31370684).

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