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Assessing the possibilities of intercropping oil palm and
pepper, under the double-row avenue system
MSc Thesis Plant Production System
Adrien-François Migeon
901219572110
June 2018
Assessing the possibilities of intercropping oil palm and
pepper, under the double-row avenue system
MSc Thesis Plant Production System
Disclaimer: this thesis report is part of an education program and hence might still contain (minor)
inaccuracies and errors.
Correct citation: Adrien-Francois Migeon, 2018, Assessing the possibilities of intercropping oil palm and
pepper, MSc Thesis Wageningen University, 33p.
Contact [email protected]
Name Student: Adrien-François Migeon
Registration number: 901219572110
Study: MSc Plant Sciences – Natural Resource Management
Chair group: Plant Production System
Code Number: PPS-80424
Supervisors: Maja Slingerlang
Meine van Noordwijk
Ni’Matul Khasanah
Examiner: Katrien Descheemaeker
Date: June, 2018
Abstract
Palm oil is the most produced oil in the world. The greatest producer is Indonesia, where 41% of the total
production is done by smallholders. More than 70% of the Indonesian smallholders rely on the palm oil
production, making them extremely vulnerable to market price changes. Of these smallholders, a great
proportion is reaching the aging phase of their plantations. Therefore, a new planting campaign will start
soon. It is the opportunity for change of paradigm. Intercropping is one of the solution mean of sustainable
intensification. The MPOB developed a new planting pattern called the Double-Row Avenue System. In this
research we look at the possibilities of intercropping oil palm and pepper (Piper nigrum) using the
WaNuLCAS model. The results show an average decrease of 3.5Mg FFB ha-1 year-1 when comparing the
conventional planting pattern and the intercropping pattern. A further decrease of 2.5Mg FFB ha-1 year-1
is observed when pepper is added to the system. On the other hand, pepper produces an additional average
of 1.9 Mg ha-1 year-1.
Table of contents
1. Introduction ................................................................................................................... 1
1.1. Sustainable production ................................................................................................. 1
1.2. Farming description ...................................................................................................... 1
1.3. Yield gap with smallholders ........................................................................................... 2
1.4. Permanent intercropping in oil palm ............................................................................... 3
1.5. Black pepper ............................................................................................................... 5
1.5.1. Plant description ...................................................................................................... 5
1.5.2. Agronomy ............................................................................................................... 5
1.5.3. Ecology ................................................................................................................... 5
1.6. Research objective ....................................................................................................... 6
2. Material and Methods ....................................................................................................... 9
2.1. Model site parameterisation ......................................................................................... 10
2.1.1. Soil sampling protocol ............................................................................................. 10
2.1.2. Climate parameters ................................................................................................ 10
2.2. Pepper parameterisation ............................................................................................. 11
2.2.1. Pepper parameters labelling protocol ......................................................................... 11
2.2.2. Pepper parameters sampling size .............................................................................. 11
2.2.3. Pepper parameters sampling protocol ........................................................................ 12
2.2.4. Biomass allocation .................................................................................................. 14
2.2.5. Tree parameterisation ............................................................................................. 14
1.7. Model validation ........................................................................................................ 14
3. Results ........................................................................................................................ 15
3.1. Pepper allometry and biomass distribution ..................................................................... 15
3.1.1. Leaf biomass ......................................................................................................... 15
3.1.2. Plagiotropic branches biomass .................................................................................. 16
3.1.3. Orthotropic branches biomass .................................................................................. 17
3.1.4. Fruit biomass ......................................................................................................... 18
3.1.5. Biomass distribution ............................................................................................... 19
3.1.6. Identification of farm effect ...................................................................................... 19
3.2. WaNuLCAS modelling results ....................................................................................... 20
3.2.1. Oil palm in monoculture .......................................................................................... 20
3.2.2. Pepper in monoculture ............................................................................................ 21
3.2.3. Oil palm and pepper under the double-row avenue system ........................................... 21
4. Discussion .................................................................................................................... 23
5. Conclusion ................................................................................................................... 25
Reference ............................................................................................................................... 27
Appendix 1 ............................................................................................................................. 29
Appendix 2 ............................................................................................................................. 31
Appendix 3 ............................................................................................................................. 31
Appendix 4 ............................................................................................................................. 32
Appendix 5 ............................................................................................................................. 32
Wageningen University and Research | Introduction 1
1. Introduction Palm oil is found in many of our daily used products, going from food products to cooking oils, to cosmetics
and bioenergy. It is the most produced oil (Woittiez et al., 2017) and the demand for palm oil is
continuously increasing. Oil palm has the highest vegetable oil yield per unit area of all oil crop (Corley et
al., 2015). Indonesia and Malaysia are the world largest producers of oil palm, with a production of 46%
and 34% of the world’s total production respectively (FaoStat, 2016). The fast increase in palm oil demand
has raised concerns. On the one hand, the increasing demand of oil leads to an increase in land-use
changes, putting at risk forests and high diversity environments. On the other hand, the palm oil production
plays an important role in the economic development of the palm oil producing countries and their rural
development.
1.1. Sustainable production The World Commission on Environment and Development of 1987 established the following definition of
sustainable development; “... development that meets the needs of the present without compromising the
ability of future generations to meet their own needs”. The delegate of the Earth Summit of Rio de Janeiro
added the concepts of “economic growth” (profit), “environmental protection” (planet) and “social equity”
(people) to the definition of sustainable development established in 1987 (Molenaar et al., 2013). This
implies that oil palm cultivations held by smallholders need to meet three characteristics to be considered
sustainable;(1) to have a long-term economic and financial viability, (2) to be responsible of the individuals
linked to the plantations and (3) to act in an environmental responsible way and aim for a conservation of
the biodiversity (Molenaar et al., 2013).
In order to meet the first characteristic (economic viability) and the third characteristic (environment
aspect), the optimisation of yield can be used thus, reducing the pressure on land-use changes. Indeed,
according to Mielke, I.S.T.A., 2010, oil palm has the highest oil yield per hectare compared to other oil
crops such as soybean, rapeseed, and sunflower, producing five to ten times more than its nearest rivals
(soybean and rapeseed) and therefore requires less land to produce the same amount of oil (Mielke,
I.S.T.A., 2010). However, with increasing demand for oil, the expansion of the oil palm production area is
inevitable. Indeed, the Indonesian ministry of agriculture reported that over 10 million hectares of land
use was dedicated to the crude palm oil production in 2014 and estimated an increase of more than 8% of
the area used for crude palm oil from the year 2014 to 2016 (Indonesian Ministry of Agriculture, 2015).
Such an expansion puts directly forests and areas with high biodiversity ecosystems at risk and creating
potential social conflicts between indigenous communities and farmers (Molenaar et al., 2013). Therefore,
optimising yields per hectare is an important element to reach sustainable farming. However, with the
increasing yields per hectare and a possible increasing income to the farmers, an additional motivation to
expand the production areas by the farmers might be created. Therefore, the reduction of pressure on land
by increasing yields can only be achieved if it is followed by adequate farming practices, agricultural
policies, and land conservation policies. In 2005, the Roundtable on Sustainable Palm Oil (RSPO) was
created to promote the sustainable palm oil production. To produce Certified Sustainable Palm Oil,
companies must comply with a series of criteria related to the protection of the environment and wildlife
but also with respect to the social, communities and workers’ interests.
1.2. Farming description The Indonesian crude palm oil production system is subdivided into three different farming categories; the
smallholders, the governmental estates, and the private estates. The smallholders represent 41% of the
land used for the palm oil production (Indonesian Ministry of Agriculture, 2015) and involve 1.7 million
smallholder households ((Indonesian Ministry of Agriculture, 2011). The Roundtable on Sustainable Palm
Oil (RSPO) defined smallholders as: “farmers growing oil palm, sometimes along with subsistence
production of other crops, where the family provides the majority of labour and the farm provides the
principal source of income and where the planted area of oil palm is usually below 25 hectares in size”
(Molenaar et al., 2013) and are characterised, in Indonesia, by an average oil palm plantation size of 3ha
(Molenaar et al., 2013). The smallholders can be classified into three subcategories; (1) the tied
smallholders, contracted to a plantation company which provides technical assistance and marketing and
supply their production to the company’s palm oil mill, (2) the independent smallholders, not bound to a
plantation company and (3) the tied + independent smallholders, who own both plantation plots tied to a
plantation company, and independent plots (Molenaar et al., 2013).
Monocropping is the main oil palm production system in all three farming categories (smallholders, private
estates, and governmental estates). The oil palm plantations are based on an equilateral triangular planting
pattern (Figure 3). This pattern is due to two main reasons; the fact that palm canopies are considered
hemispherical, an equilateral triangular pattern allows a better land use than a square pattern and
secondly, the fact that palms are monocotyledon plants, therefore, do not grow irregularly to fill in the
open spaces like dicotyledonous plants would do (Corley et al., 2003).
Wageningen University and Research | Introduction 2
1.3. Yield gap with smallholders
According to Goh et al., (1994), there is a fairly consistent correlation in the yield pattern (ton FFB ha-1
year-1) related to the age of the oil palm (Figure 1). From the data analysed by Goh et al., (1994), it seems
that there is a yield decline linked to the age of the palm. This yield decline is estimated to be at 20 years
of age. The decrease in yield is due to two main reasons; the physiological decline of the plant creating a
reduction of the yield and in addition to this, the harvesting difficulties encountered due to the increasing
tree height.
Figure 1, Representation of the distribution of yield with age with the different productions phases. (After Goh et al., 1994 and
Ling, 2012)
Indonesian tied smallholders have an average palm age of 19 years and therefore will require a replanting
of the plot in the near future. On average, peak productions are reached between the age 9 to 18 years
and gradually decline after that (Figure 1). However, palms can be kept for production for up to 30 years
(United States Department of Agriculture, 2012). In addition to this, one fifth of the independent
smallholders may require replanting due to poor palm varieties (Molenaar et al., 2013).
In production ecology, based on oil palm literature, three different production levels can be distinguished;
(1) the potential yield, (2) the site yield potential and (3) the actual yield.
(1) The potential yield is determined by yield-defining factors; photosynthetically active radiations,
temperature, ambient CO2 concentrations and crop genetic characteristics (van Ittersum et al., 1997).
Evans, (1993) defines the potential yield as the yield of a cultivar, when grown in environments to which
it is adapted; with nutrients and water non-limiting; and with pests, diseases, weeds lodging, and other
stresses effectively controlled.
(2) The site yield potential is defined as the yield obtained on a specific site, with natural water supply,
nutrients supplied at optimum rates, and agronomic and disease control measures implemented to a high
standard (Corley et al., 2015). The site yield potential includes management decisions such as planting
material and density (Woittiez et al., 2016).
(3) The actual yield is determined by yield reducing factors; weeds, pests and diseases (van Ittersum
et al., 1997).
Under the current agricultural practices, the actual yields of FFB (t ha-1 year-1) for Indonesia and Malaysia
are 17 and 21 respectively and the potential yields are of about 36 and 28 respectively (Table 1).
Therefore, the foreseen upcoming replanting of the plantation may create an opportunity to close down
the yield gap and, thus, improve yields.
Table 1, Fresh fruit bunch (FFB) and Crude Palm Oil (CPO) yields comparison between actual yield and potential yield (After Woittiez et al., 2016)
Country Fresh Fruit Bunch (t ha-1 year-1) Crude Palm Oil (t ha-1 year-1)
Actual Yield Potential Yield Actual Yield Potential Yield
Indonesia 17 32 - 40 3.8 8 - 10
Malaysia 21 24 - 32 4.2 6 - 8
Wageningen University and Research | Introduction 3
1.4. Permanent intercropping in oil palm
Despite the needs of replanting an ageing plot, the financial aspect of replanting is a major drawback for
the farmer. For smallholders, the replanting stage is a very costly practice. Various factors such as the
investment needed for replanting, the lack of income during the immature stage, the low yield in the early
stage of production and the long-term payback of the investment make the replanting of the plots very
challenging for the farmers. It is reported that 35% of the farmers save money for replanting on a regular
basis and 28% occasionally (Molenaar et al., 2013). However, they do not believe that their savings are
sufficient to cover the full financial needs during replantation. The effect of the investment needed for
replanting will differ with the importance of the income brought in the household through the production
of palm oil. It is known that 18% of the smallholders fully rely on oil palm as the main income and for 69%
of the farmers, it represents more than half of their income, representing over 300.000 and 1.100.000
farmers respectively.
This financial burden induced by the replanting process can, however, be diminished by the use of
alternative strategies such as intercropping. Intercropping is not interesting to the big plantations due to
managerial difficulties (Corley et al., 2003). However, on average smallholders have plots of around 2ha,
it may be an interesting solution to them. Indeed, the use of the additive intercropping design may bring
an income during the immature phase of the oil palm and an additional income during the production
phases of the oil palm in the case of a cash crop (Nuertey et al., 2000), or food in the case of a subsistence
crop. In addition to this, intercropping also helps in reducing the cost of weeding as well as maximising the
land use and therefore limit the pressure exerted by farmers on forest areas and high biodiversity
ecosystems (Nchanji et al., 2015). Okyere et al., (2014) and Hartley, (1988), state that according to their
research, in the case of a previous intercropping during the immature phase of the oil palm, the
intercropping has no adverse residual effect on the growth, development and yield of the oil palm during
the following production phases. This suggests that under good management practices, the food crop
previously intercropped does not influence the nutrient uptake of the oil palm.
Despite the many advantages brought by intercropping oil palm with cash or subsistence crops, the
intercropping stage can only be maintained for the beginning of the immature phase of the oil palm.
Indeed, under a conventional triangular oil palm planting system (Figure 2a), certain crops can be
integrated for a period of 1.5 to 2 years after the establishment of the plantation. Beyond that period of
time, it becomes less suitable for intercropping as the fronds of the palm start to encroach the surrounding
palms (Suboh et al, 2009). This leads to a limitation of light penetration through the canopy (Figure 3).
Figure 2, Equilateral triangular pattern (A), double row avenue planting pattern (B) (After Suboh et al., 2009)
However, in order to reduce the effect of light limitation induced by the intercropping of oil palm under the
conventional equilateral triangular pattern (Figure 2a), the Malaysian Palm Oil Board (MPOB) has developed
a different intercropping pattern called “double row avenue system” (Figure 2b). This plantation pattern
allows the integration of other crops in the avenue between the rows of oil palm. Indeed, Mohd Haniff et
al., (2003), measured that the photosynthetically active radiation (PAR) penetrating through the canopy
was increased by as much as 50% in the double-row avenue as compared to the conventional triangular
planting pattern. He suggests that the open space on the avenues between the oil palm rows is suitable
for establishing a crop as the mean PAR transmitted through the canopy was 40%. This change in PAR was
correlated to the leaf area index of the two systems, as it was higher under the conventional planting
pattern.
Wageningen University and Research | Introduction 4
Figure 3, Distribution of the photosynthetically active radiation penetrating the oil palm canopy reaching the soil measured along
a transect between two palms, for the conventional triangular pattern (dashed line) and double-row avenue (solid line) (After
Mohd Haniff et al., 2003 and Suboh et al., 2009)
With the use of the double-row system, the density is of 138 trees ha-1 on the overall system which is a
similar value compared to the conventional triangular system, with a density 139 trees ha-1. However, the
density within the row is of 192 trees ha-1. According to Goh, (1982), there is a decrease in yield of FFB
palm-1 corresponding to an increase of the planting density, therefore a lower yield would be expected in
the double-row avenue system (Figure 4). However, according to the results from the various experiments
conducted by the MPOB, it appears that the yield of FFB under the double-row avenue system is similar to
the yield in the conventional planting pattern (Table 2). This response in yield may be due to the effect of
the planting density within the palm rows and to the additional PAR made available by the border effect of
the canopy adjacent to the avenue. No studies have been found on this matter.
Figure 4, Yield of Fresh Fruit Bunch per palm according to the planting density (Goh, 1982)
Table 2, Average FFB yields (t ha-1 year-1) in the first four years of harvest at Maah Plantation, KLIA, Sepang, Malaysia (After Suboh et al., 2009)
Oil palm Planting System
FFB Yield (t ha-1 year-1)
1st year harvest
2nd year harvest
3rd year harvest
4th year harvest
Conventional triangular Control plot 3.52 9.04 24.66 26.31 Double-row avenue Integrated with hill paddy (Oryza sativa) 3.21 10.71 23.25 23.15 Integrated with sorghum (Sorghum bicolor) 4.07 11.67 25.40 22.96 Integrated with tongkat ali (Eurycoma longifolia) 3.48 9.08 21.97 22.10 Integrated with pigeon pea (Cajanus cajan) 4.21 9.73 23.20 25.19 Control plot 3.98 9.77 27.12 23.36
The integration of crops with oil palm will optimise the land-use and provide additional income to the farmer
or subsistence food. Table 2 shows that several crops are suitable to be integrated with the oil palm under
the double-row avenue system.
0
10
20
30
40
50
60
70
80
90
100
0 2 4 6 8 10 12 14 16
Ph
oto
syn
thec
ally
Act
ive
Rad
iati
on
(%
)
Distance between palms (m)
Wageningen University and Research | Introduction 5
1.5. Black pepper
Black pepper (Piper nigrum L.), also known as the King of Spices is the most important and widely used
spice in the world. It originates from the tropical forests of Western Ghats, South India and was taken
later, due to European colonisation of the 17th century, to Indonesia and Malaysia. It is now produced in
over 26 countries, around the tropical belt. In Malaysia and Indonesia, it is considered a smallholder’s crop,
with an average holding of 0.1 to 0.4 ha per family (Ravindran et al., 2003).
The product consumed is a dried drupe that grows on a spike and is harvested before ripening. White
pepper is from the same plant, it is the post-harvest process that differs. Indeed, the fruits are collected
when the drupe is ripe (red or orange colour), the pericarp is removed and the seeds are dried (Ravindran
et al., 2003).
1.5.1. Plant description
Black Pepper is a perennial, woody and climbing plant from the family of the Piperaceae. Under farming
conditions, the adult plant grows on a support. Pepper does not usually have more than 10 main roots.
The feeder roots are distributed in the first 30–40cm of the soil. The water absorbing roots, however,
penetrate the soil to a depth of up to 4m (Terada et al., 1971). Independently of the age, most roots (67-
85%) are found on the first 0-30 cm of the soil depth and at 35-70cm of the base of the plant (Ramos et
al., 1984). The pepper plant has two distinct types of branches, the orthotropic branches and the
plagiotropic branches, where each has their own functions. The orthotropic branches are the climbing
branches. They remain vegetative and have short adventitious roots present at the nodes, allowing
adherence to the support on which it grows. The plagiotropic branches, on the other hand, have the ability
to become generative. They produce higher-order branches as well as inflorescences (Guigaz, 2002).
Inflorescences are spike shaped and appear opposite to the leaves on plagiotropic branches. They hold 50
to 150 flowers on each spike. The flowers are usually bisexual. Geitonogamy is believed to be the common
mode of autogamous pollination (Corley et al., 2003). Geitonogamy is a self-pollination mechanism by
which the pollen descends due to the effect of gravity combined with the rain water or dew drops. This is
an efficient mechanism amongst plants with long pendent inflorescences. However, heavy rain has a
negative effect on pollination (Ananda, 1924). After fertilisation, the ovary takes 8 to 9 months to mature.
The leaf area per plant in bush pepper is primarily dependant on the increase of leaf area per leaf whereas
for vine pepper, the leaf area per plant is dependent on the number of leaves produced. There is a
significant difference in the biomass distribution, the number of leaves and, the total leaf area, between
vine pepper and bush pepper (Geetha et al., 1992).
The productivity of a plant depends on its efficiency in harvesting solar energy for the metabolite production
and its partitioning efficiency of the metabolites produced (Bindra et al., 1977). It has been shown for
black pepper that a higher light availability before flowering induced a greater leaf area and a more compact
canopy structure with shorter lateral shoots. This leads to a higher accumulation of metabolites, more
spikes, more flowers and therefore, a greater number of fruit and a higher dry matter of fruits per plant
(Mathai et al., 1988). Mathai et al., (1990) observed a significant relationship between biomass production
and economic yield. However, when illumination exceeds 900 µmol s-1 m-2, the photosynthetic rate
decreases (Mathai, 1983).
1.5.2. Agronomy
Even though black pepper produces drupes, the commercial propagation is only done through vegetative
propagation, with the use of cuttings made from orthotropic shoots. Cuttings are usually kept in a nursery
until transplanting. One or two months after transplanting, the plant becomes vigorous. Posts have to be
installed on the field to the required distance before transplanting.
Pepper can be harvested from April-June until August-September in South-East Asia. Harvesting is done
every two weeks. Black pepper plantations have an economic lifespan of 15 to 20 years.
1.5.3. Ecology
Black pepper is suited for the tropical climate with a well distributed annual rainfall of 2000-4000mm and
with a temperature of 25-30°C and a relative humidity of 65-95%. The crop grows best below 500m on
the equator but can be grown at altitudes of up to 1500m. It grows on soils ranging from heavy clay to
light sandy clays. The soil needs to be deep, well drained but with a sufficient water-holding capacity for
the plant to deal with water stress (Westphal et al., 1989).
Wageningen University and Research | Introduction 6
1.6. Research objective
Raja Zulkifli et al., (2016) conducted a research on the integration of black pepper (Piper nigrum) under
the double-row avenue system, in the MPOB research stations of Belega located in the state of Sarawak
and Keratong in the state of Pahang, Malaysia. They concluded that black pepper is a suitable crop to be
integrated with oil palm under the double-row avenue system as it is technically and economically feasible
to be practised by the smallholders. In addition to this, it is stated that the integration of crop with oil palm
optimises land use and provides and an additional long-term income to the farmer. However, in comparison
to the results found by Suboh et al., (2009), the measured yield is lower in the 3rd and 4th year of harvest
(Table 3). This decrease in yield is not explained. The results obtained by Raja Zulkifli et al., (2016) have
been compared to the MPOB oil palm production statistics at the Malaysian national level and to the
corresponding regional level (Pahang and Sarawak) however, no comparable decreasing trends in yields
have been found.
Table 3, Comparison of the FFB yields (t ha-1 year-1) in the first four years of harvest obtained by Suboh et al., (2009) and Raja Zulkifli et al., (2016).
Reference Oil palm Planting System FFB Yield (t ha-1 year-1)
1st year harvest
2nd year harvest
3rd year harvest
4th year harvest
Suboh et al., (2009) Conventional triangular
Control plot 3.52 9.04 24.66 26.31
Double-row avenue
Mean value of integrated crops 3.74 10.29 23.45 23.35
Control plot 3.98 9.77 27.12 23.36
Raja Zulkifli et al., (2016) Conventional triangular
Control plot n.a n.a 15.10 13.60*
Double-row avenue
Integrated with black pepper 6.00 13.00 16.90 13.20*
Control plot 5.50 10.30 14.40 13.00*
Note: * Nine months record, we assume a yield of 18.13 t ha-1 year-1 for the conventional triangular control plot, 17.6 t ha-1 year-1 for the double-row avenue system integrated with black pepper and 17.3 t ha-1 year-1 for the double-row avenue system control plot ; n.a = not available
Intercropping is of little interest to the large plantations. Therefore, there is a scarce amount of literature
available on this matter (Corley et al., 2003). There is little research on oil palm intercropped with black
pepper. The use of model simulations to investigate the potentialities and restrictions of the intercropping
of oil palm and black pepper is an important step in order to evaluate the possibilities of the system in a
less costly and time-consuming way. The use of model simulation would allow exploring the system in the
future and, to identify its limiting factors and management strategies (Pinto et al., 2005)
The WaNulCAS model (Water, Nutrient and Light Capture in Agroforestry Systems) is a tree-soil-crop
interaction model used for agroforestry systems. It allows the assessment of the water, nutrients and light
interactions in the system. The model represents a four-layer soil profile (vertical), with four spatial zones
(horizontal) (Figure 5) and allows a description of the uptake of water and nutrients on the basis of root
length densities of both plants grown on the plot (van Noordwijk et al., 1999). The WaNulCAS model
simulates dynamic processes in a spatial and field scale environment, such as the simulation of above and
below ground plant growth within a dynamic biophysical environment (Walker et al., 2007).
The vegetative and generative duration of the crops chosen is used as an input. Each crop should be
specified on the basis of its maximum dry matter production rate per day (kg DM m-2 day-1) and on its
relative light use efficiency (dry matter produced per unit of light intercepted) and its leaf weight ratio as
a function of the crop stage (van Noordwijk et al., 1999).
The model incorporates management options such as tree spacing, pruning and choice of species (van
Noordwijk et al., 1999).
Wageningen University and Research | Introduction 7
Figure 5, General layout of the zones and layers in the Wanulcas model (After van Noordwijk et al., 1999)
The interactions between the species selected for the modelling are characterised by many parameters.
These parameters are defined under a crop library in the case of a crop or a tree library in the case of a
tree. The crop library holds a database for specific crop parameters and crop related input-output for the
system simulated. Overall, there are 58 inputs parameters (Appendix 1) including 5 growth parameters as
a function of crop stage (van Noordwijk et al., 2011). However, the crop library for black pepper does not
exist.
The aims of the study are twofold:
1. Develop allometric models for pepper, to represent quantitative relations between the growth of
different organs on the plant.
2. Aim at the creation of a crop library for pepper, using the previously created allometric models.
a. First, integrate the allometric models and biomass distribution equation in the WaNuLCAS
model.
b. Second, parametrisation and calibration of the WaNuLCAS model
The aim of this study is the creation of a crop library for black pepper, enabling the use of the WaNuLCAS
model to research the growth and production of oil palm and black pepper (Piper nigrum) under the double-
row avenue system, in the MPOB research station in Belega, Malaysia. This will allow to answer the research
questions:
R.Q.1: Is the production of oil palm and black pepper possible in the long term under the double-row
avenue system?
R.Q. 2: Does the oil palm and black pepper use resources from different niches? Is there a competition
for resources such as light, nutrients and water?
Wageningen University and Research | 8
Wageningen University and Research | Material and Methods 9
2. Material and Methods
Raja Zulkifli et al., (2016) conducted an experiment measuring the yields obtained for both oil palm and
black pepper. However, it only lasted for the time of four years after the first harvest of oil palm. The
objective of this study is to use the WaNuLCAS Model to investigate, in the long term, the biophysical
interactions and the performance of oil palm and black pepper under the double-row avenue system.
The experiment consists of twofold; the creation of allometric equations allowing the prediction of the
growth and yield of black pepper, and the simulation of intercropping black pepper and oil palm under the
double-row avenue system. The experimental phase was conducted in four distinctive steps; (1) a model
site parameterisation was performed. This involves a site parameterisation describing the abiotic conditions
of the investigation site (rainfall, temperature, soil description) and the definition of the four spatial zones
with their attributed ground cover; (2) a crop parameterisation and the creation of a crop library for the
black pepper, including the creation of allometric equations; (3) a model simulation was carried out using
WaNuLCAS to predict the growth of pepper and oil palm under the double-row avenue system; (4) a model
validation was performed, based on a comparative data analysis between the model predicted data and
the observed data on the MPOB research station, in Belaga.
In order to collect the data allowing the creation of allometric equations of black pepper and the creation
of a pepper model used in the crop library, a partnership was established with the World Agroforestry
Center (ICRAF), in Bogor, Indonesia. This allowed us to collaborate with three local farmers and access
pepper fields in the region of Konawe, Southeast Sulawesi, Indonesia (Figure 6). Interviews were
performed with farmers. This allowed us to choose the farmers with whom we would collaborate, based on
the farming practices, the pepper variety used, the maintenance of the plantation, the fertilizer application
and the presence of disease (Table 4).
Table 4, Agricultural practices description for the three farmers
Farming Practice Farmer Mr. Muksin Mr. Ramli Haji Tonga
Pepper variety Local Local Local
Propagation Orthotropic Branch Cuttings Orthotropic Branch Cuttings Orthotropic Branch Cuttings
Pepper support Glyricidia Glyricidia Glyricidia
Planting distance 2.5m*2.5m 2.5m*2.5m 2.5m*2.5m
Fertiliser Application Chicken manure + NPK NPK Chicken manure
Presence of diseases No Occasional borer No
Figure 6, Study zone where the three farms are located. The inset map on the left-top corner shows in the grey frame the
location of the region of Sulawesi in Indonesia. The greyed zone on the map shows the administrative area of Konawe.
Wageningen University and Research | Material and Methods 10
2.1. Model site parameterisation
In the WaNuLCAS Model, four zones spatial zones need to be defined, where the ground cover and the
zone width needs to be defined. Moreover, the model has a mirror effect at each of the borders of the four
zones. Thus, the defined zones are repeated oppositely to the left of zone 1 and to the right of zone 4
(Figure 7). The horizontal zones (Figure 7) were established in the model according to the double-row
avenue system pattern established by Suboh et al., (2009) (Figure 2). Indeed, a distance of 8.5m between
the palms within the palm row and a distance of 15.2m of open avenue are respected. Moreover, a distance
of 2.5m between the pepper plants is established. The soil layer depth has to be set accordingly to the
rooting depth of black pepper and oil palm. The palm density is set at 138 trees ha-1.
Figure 7, the double row-avenue system configuration as used in the WaNuLCAS model. The oil palm tree is located on the right
of zone 1. Zone 2 is left empty, as the palm tree will grow and create shade within the open avenue. Zone 3 has a pepper plant
implemented in its middle. Zone 4 has a pepper plant implemented to its right. The model is parameterised in order to reduce
the production of pepper in zone 4 by 50 percent, thus allowing the simulation of a full pepper plant when taking in consideration
the mirror effect occurring when using the WaNuLCAS model.
2.1.1. Soil sampling protocol
In order to determine the soil characteristics (Table 5), a soil sampling and analysis was carried out. A
random sampling was carried out across the plots in order to cover soil variability. Three samples per
depth, per farm, for the three farms were collected. The depth of the sample collection was 0 to 20cm, 20
to 40 cm and 40 to 60cm. The samples were collected using a soil auger. In order to keep track of the
sampling locations, the coordinates were recorded using a handheld GPS receiver (Garmin Montana 600).
Once the samples collected, they were bulked together, according to the soil depth, and thoroughly mixed.
Sub-samples were collected into smaller batches to match the recommended amount of soil needed by the
laboratory conducting the soil analysis.
Table 5, Soil Analysis for soil site parameterisation
Horizons Texture pH Nutrient Content & Cation Exchange Capacity
Soil Depth Sand Silt Clay H2O KCl C organic P2O5 CEC
cm % % % [] [] % ppm Cmolc kg-1
0-20 13 54 33 4.5 3.7 1.13 15.3 13.20
20-40 14 55 31 4.7 3.8 0.75 4.9 12.43
40-60 12 61 27 4.6 3.7 0.56 4.9 12.38
2.1.2. Climate parameters
The climate parameters (Table 6) used in the model were based on the weather data gathered by the local
weather station.
Table 6, Data required for site parameterisation
Climate
Parameters Unit Source
Rainfall (daily or monthly) mm Weather station data
Daily Soil temperature °C Weather station Data
Wageningen University and Research | Material and Methods 11
2.2. Pepper parameterisation
In order to simulate the phenological development of black pepper in the model, a black pepper Crop
Library was established. Given the fact that the crop library comprises 58 different input parameters
(Appendix 1). The crop library is partly based on field data collected in three black pepper plantations
(Table 7), in the region of Konawe, Southeast Sulawesi, Indonesia, as well as through information found
in literature and defaults values set in the WaNuLCAS model (M. van Noordwijk, personal communication,
8 June 2017).
Table 7, Black pepper parameters collected in monocultures
Parameters unit Acronym
Ratio of height increment to biomass increment [] Cq_HBiomConv Production of dry matter kg m-2 day-1 Cq_GroMax Length of the vegetative stage days Cq_TimeVeg Length of the generative stage days Cq_TimeGen Maximum Leaf Area [] Cq_LAIMax
Relative light intensity at which shading starts to affect the crop growth [] Cq_RelLightMaxGr
The WaNuLCAS model includes a Crop Management section where all the different agricultural practices
conducted through the experiment are referenced in the model. Therefore, in addition to the physiological
characteristics needed to run the model, managerial information was provided by the farmers, allowing
the model to consider those various agricultural practices during the results computation. These practices
include; timing of planting, timing of fertiliser and organic matter application and harvesting of crop
biomass.
2.2.1. Pepper parameters labelling protocol
The measurements and samples were classified according to each plant thus, assigned with the plant
number and the farm it was located on. In addition to this, each plant was subdivided into sections of 50cm
high. The number of sections present per plant was based on the plant height (Figure 8). The sampling
classification method was used for all material sampled.
Figure 8, Representation of a black pepper plant growing on a pole.
The plant is divided into sections set according to the height.
2.2.2. Pepper parameters sampling size
During the pepper data collection, we decided to sample 30 plants non-destructively. We measured 10
pepper plants in three different farms with similar agricultural practices (Table 4). We opted for the sample
size of 10 plants per farm on the basis of expecting to have homogeneous data within each farm. It was
decided to collect data from three different farms as this would enable us to detect a farm effect in the
data in case of its presence. Moreover, due to time limitations, the above-mentioned sample size seemed
the most feasible. A total of more than 15.000 measurements were performed for the plagiotropic branches
alone.
In addition to this, 2 plants were used for destructive measurement in order to create volume conversion
coefficients and density coefficients.
Wageningen University and Research | Material and Methods 12
2.2.3. Pepper parameters sampling protocol
For the non-destructive measurement of the 30 plants, a total of 6 general parameters were collected per
plant; (1) Plant height, (2) Plant DBH, (3) Number of intersections between the orthotropic branches and
a vertical axis following the plant support, (4) the diameter of each orthotropic branch at each section (0,
50, 100, 150, 200cm), (5) the number of leaves per section, and (6) the number of generative organs per
section, according to their development stage. Moreover, 5 plagiotropic branches were analysed per section
per plant. The measurements consist of measuring the length and diameter of each internode and recording
its position in the branch structure. In addition to this, each leaf and generative organ supported on the
measured branch is recorded according to its position on the branch. Finally, 5 leaves per section per plant
were collected in order to establish leaf area and leaf biomass.
2.2.3.1. General parameters
1) Plant height
The plant height is determined by the distance between the ground and the highest leaf observed. The
pepper plant heights were measured using a plumb line and a measuring tape. The plumb line was attached
to a rod with a spirit level to ensure the horizontality of the rod when placing it at the top of the plant to
do the height measurement. The plumb line was allowed to slide through the rod allowing its adjustment
to the ground level. Sections of 50 cm were created allowing for classification of the samples. The highest
section was determined based on the highest multiple of 50cm below the highest measured leaf.
2) Plant diameter
The diameter of the canopy of the pepper plant was measured at breast height. In order to determine the
diameter, the circumference of the plant was measured using a flexible measuring tape.
3) Leaf area
The leaf area is based on the average individual leaf area observed per section. The leaf area is
characterised by the one-sided area of a fresh leaf. Five leaves per section were sampled, thus the leaf
area of 745 leaves was measured.
Each leaf included the leaf blade and its petiole. An image of each leaf was taken and analysed with an
image analysis software. The leaves were placed under a transparent plexiglass cover allowing a flat
positioning of the leaves before the capture of the image and were placed on a white background. The
measurements were obtained using an image analysis software, ImageJ, with the image processing
package FIJI. The software measures digital images by using the RGB (Red, Green and Blue) value of each
pixel to identify the leaf and scale regions in each image (Appendix 2). The conversion factor between the
pixel size and the true size (cm) represented by each pixel was set by establishing a scale. The imaging of
each leaf was done using a digital camera (Olympus OMD-EM5) placed on a tripod with the centre-column
installed upside down.
4) Leaf Biomass
The total number of leaves per section was recorded. In addition to this, for each plant, 5 leaves were
sampled per section. Each sampled leaf was analysed. The fresh biomass and dry biomass was measured.
To determine the individual leaf biomass (kg DM), the leaf samples were placed in a drying oven at 70°C
for a minimum time of 72 hours. Once the leaves dry, they were removed from the oven and their biomass
was determined by measuring the dry matter of each leaf using an electronic scale with a resolution of
0.01g. The dry matter biomass of 745 leaves was established.
Based on these measurements, we can establish the leaf biomass per section of 50cm, thus, if the height
of the plant is known, the total leaf biomass per plant (equation 1).
(1) 𝑇𝑜𝑡𝑎𝑙 𝐿𝑒𝑎𝑓 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 = ∑ (𝑛 𝑙𝑒𝑎𝑣𝑒𝑠 𝑝𝑒𝑟 𝑠𝑒𝑐𝑡𝑖𝑜𝑛 ∗ 𝐴𝑣𝑒𝑟𝑎𝑔𝑒 𝐿𝑒𝑎𝑓 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 𝑝𝑒𝑟 𝑠𝑒𝑐𝑡𝑖𝑜𝑛 (𝑔))
𝑛 𝑠𝑒𝑐𝑡𝑖𝑜𝑛𝑠
𝑖=0
2.2.3.2. Plagiotropic branch biomass
The plagiotropic branches for the 30 plants measured non-destructively were measured. For each plant, 5
branches were measured per each section of 50cm high. Each of the internodes composing these branches
were measured, based on their length and diameter, in order to establish a Theoretical Volume for each
internode (equation 2). The lengths and diameters were measured with an electronic digital calliper with a
resolution of 0.01 mm.
Wageningen University and Research | Material and Methods 13
(2) 𝑇ℎ𝑒𝑟𝑜𝑟𝑒𝑡𝑖𝑐𝑎𝑙 𝐵𝑟𝑎𝑛𝑐ℎ 𝑉𝑜𝑙𝑢𝑚𝑒 (𝑐𝑚3) = (𝐿𝑒𝑛𝑔𝑡ℎ ∗ 𝐷𝑖𝑎𝑚𝑒𝑡𝑒𝑟2 ∗ 𝜋
4) ∗ 10−3
In addition to this, the structure of the branch was recorded. Moreover, the number of leaves at their given
position was recorded for the plagiotropic branches.
In order to convert the Theoretical Volume of the plagiotropic branches into their Biomass, we needed to
determine two transformation coefficients allowing to estimate the Real Branch Volume and the Branch
Biomass. The plagiotropic branches collected during the destructive sampling of the 2 pepper plants were
used to create the two transformation coefficients. In total, 737 plagiotropic internodes, originating from
the destructive measurement, were analysed. Their
length, diameter, fresh biomass, dry biomass and volume
were recorded. The length and diameter were measured
with an electronic digital calliper with a resolution of 0.01
mm. All weight measurements were carried out using an
electronic scale with a resolution of 0.01g. The fresh
biomass was measured using the electronic scale. In order
to measure the dry matter, the samples were placed in a
drying oven at a temperature of 70 degree Celsius for a
minimum of 72 hours and weighted using an electronic
scale. The volume of each internode, at its fresh stage, was
measured using a suspended hydrostatic measurement
technique, where the measured branch is immersed under
the surface of a liquid of known density (demineralised
water at 24 degree Celsius) in a container placed on the
electronic scale. The branch suspended in the water is
stationary, therefore we may conclude that the forces
exerted on the branch are balanced. Indeed, the gravitational (g) force is canceled out by the buyancy
force (b) and the tension force (t) (Figure 9). The branch immersed in water is equivalent to a volume of
water exactly the same size and shape. Therefore, the volume of the immersed branch is the increase in
weight divided by the density of the fluid (equation 2).
Where ρ is the density of the fluid at a given temperature, 𝛥𝑊 is the change in weight expressed on the
scale after the immersion of the branch and V is the unknown volume.
These recorded measurements allow to establish a transformation coefficient enabling the conversion of
the Theoretical Volume to the Real Volume (equation 3) and a density coefficient to transform the Real
Volume to Branch Biomass (equation 4). These two coefficients are obtained using linear regression
models, where Theoretical Volume is plotted against Real Volume and Real Volume is plotted against
Biomass. Indeed, the relationship between the dependent variables and independent variables will be
described by the regression analysis, thus allowing to estimate the values of the dependent variables
according to the independent variable.
(3) 𝑅𝑒𝑎𝑙 𝐵𝑟𝑎𝑛𝑐ℎ 𝑉𝑜𝑙𝑢𝑚𝑒 (𝑐𝑚3) = ((𝐿𝑒𝑛𝑔𝑡ℎ ∗ 𝐷𝑖𝑎𝑚𝑒𝑡𝑒𝑟2 ∗ 𝜋
4) ∗ 10−3) ∗ 𝑡𝑟𝑎𝑛𝑠𝑓𝑜𝑟𝑚𝑎𝑡𝑖𝑜𝑛 𝑐𝑜𝑒𝑓𝑓.
(4) 𝐵𝑟𝑎𝑛𝑐ℎ 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 (𝑔) = [((𝐿𝑒𝑛𝑔𝑡ℎ ∗ 𝐷𝑖𝑎𝑚𝑒𝑡𝑒𝑟2 ∗ 𝜋
4) ∗ 10−3) ∗ 𝑡𝑟𝑎𝑛𝑠𝑓𝑜𝑟𝑚𝑎𝑡𝑖𝑜𝑛 𝑐𝑜𝑒𝑓𝑓. ] ∗ 𝐷𝑒𝑛𝑠𝑖𝑡𝑦
The biomass of each of the plagiotropic branch measured non-destructively may now be assessed based
on the two transformation factors previously established. Thus, to assess the total biomass of the
plagiotropic branches per plant, the relationship between a given branch biomass and the number of leaves
it holds was analysed (equation 5). In order to analyse the relationship between each branch biomass and
the number of leaves it holds, a linear regression analysis is conducted. The number of observations used
to carry out the regression analysis is n=646.
(5) 𝑃𝑙𝑎𝑔𝑖𝑜𝑡𝑟𝑜𝑝𝑖𝑐 𝐵𝑟𝑎𝑛𝑐ℎ 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 𝑝𝑒𝑟 𝑃𝑙𝑎𝑛𝑡 = [ ∑ (𝑎𝑣𝑒𝑟𝑎𝑔𝑒 𝑛𝑢𝑚𝑏𝑒𝑟 𝑜𝑓 𝑙𝑒𝑎𝑣𝑒𝑠 𝑝𝑒𝑟 𝑠𝑒𝑐𝑡𝑖𝑜𝑛)
𝑛 𝑠𝑒𝑐𝑡𝑖𝑜𝑛𝑠
𝑖=0
] ∗ 𝐵𝑟𝑎𝑛𝑐ℎ 𝑐𝑜𝑒𝑓𝑓.
(2) 𝑉 =𝛥𝑊
𝜌
Figure 9, Hydrostatic measurement set-up.
(Hugues, 2005)
Wageningen University and Research | Material and Methods 14
2.2.3.3. Orthotropic branch biomass
The data used was recorded during the non-destructive measurement of the 30 plants. In order to
determine the volume of the orthotropic branches measured, we used the formula used to find the volume
of a conical frustum (equation 6). This equation is commonly used in forestry to establish branch volume.
The diameter of the internode of the orthotropic branches were measured at each of the intersections of
50cm, including ground level (0cm). In order to find the volume of the internodes of the orthotropic
branches at a given section, the sum of the diameters at the lower section (𝐷𝑎) and the sum of the
diameters at the upper section (𝐷𝑏) of the orthotropic branches were used.
(6) 𝑇𝑜𝑡𝑎𝑙 𝑉𝑜𝑙𝑢𝑚𝑒 𝑂𝑟𝑡ℎ𝑜𝑡𝑟𝑜𝑝𝑖𝑐 𝐵𝑟𝑎𝑛𝑐ℎ𝑒𝑠 𝑝𝑒𝑟 𝑆𝑒𝑐𝑡𝑖𝑜𝑛 =𝜋 ∗ ℎ + (𝐷𝑎
2 + 𝐷𝑏2 + 𝐷𝑎 ∗ 𝐷𝑏)
12
To determine the biomass of the sum of the orthotropic branches per section (equation 7), we used a
density coefficient that is determined using the orthotropic branches collected during the destructive
sampling. The procedure to determine the density was the same as for the plagiotropic branches.
(7) 𝑇𝑜𝑡𝑎𝑙 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 𝑂𝑟𝑡ℎ𝑜𝑡𝑟𝑜𝑝𝑖𝑐 𝐵𝑟𝑎𝑛𝑐ℎ 𝑝𝑒𝑟 𝑆𝑒𝑐𝑡𝑖𝑜𝑛 = 𝜋 ∗ ℎ + (𝐷𝑎
2 + 𝐷𝑏2 + 𝐷𝑎 ∗ 𝐷𝑏)
12∗ 𝐷𝑒𝑛𝑠𝑖𝑡𝑦
2.2.3.4. Fruit biomass
During the destructive measurement of the two pepper plants, a total of 619 generative organs were
collected. During this sampling phase, no distinction was made between the different development stages
of the generative organs; flower, semi-developed spike and fully developed spike. All the 619 samples
were analysed. The fresh biomass and dry biomass was established using an electronic scale with a
precision of 0.01g. In order to dry the samples, the generative organs were placed in a drying oven at 70
degrees Celsius for a period of 72 hours. Based on the data obtained, we were able to establish an average
generative organ biomass.
During the non-destructive measurement of the 30 plants, the number of generative organs and the
number of leaves were counted for each section. The relationship between the total number of leaves per
plant and the total number of fruits per plant allowed to estimate the fruit biomass per section, based on
the number of leaves observed. In order to analyse the relationship between the number of fruits per
section and the number of leaves per section, a linear regression analysis was conducted between the
independent variable; Number of leaves per section and the dependent variable; Number of fruits per
section. The number of observations used to carry out the regression analysis is n=124.
In order to find out the fruit biomass, we need to multiply the amount of fruits by the average biomass
(equation 8);
(8) 𝐹𝑟𝑢𝑖𝑡 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 = 𝑛𝑢𝑚𝑏𝑒𝑟 𝑜𝑓 𝑓𝑟𝑢𝑖𝑡𝑠 ∗ 𝑎𝑣𝑒𝑟𝑎𝑔𝑒 𝑓𝑟𝑢𝑖𝑡 𝑏𝑖𝑜𝑚𝑎𝑠𝑠
2.2.4. Biomass allocation
Pepper yield estimation may be now conducted through the use of allometric equations. Indeed, an
allometric equation represents the quantitative relation between the growth of two or more organs of the
same plant (Richards, 1969). It, therefore, becomes possible to assess a physiological status of a growing
plant by analysing the allometric relationship to a different organ.
The allometric equations used were derived from the different empirical parameters collected. This allowed
to establish the distribution of growth over the different plant organs (equation 9).
(9) 𝑃𝑙𝑎𝑛𝑡 𝐵𝑖𝑜𝑚𝑎𝑠𝑠 = (𝛼 𝐿𝑒𝑎𝑓 𝐵𝑖𝑜𝑚𝑎𝑠𝑠) + (𝛽 𝐹𝑟𝑢𝑖𝑡 𝐵𝑖𝑜𝑚𝑎𝑠𝑠) + (𝛿 𝑂𝑟𝑡ℎ𝑜 𝐵𝑖𝑜𝑚𝑎𝑠𝑠) + (𝜖 𝑃𝑙𝑎𝑔𝑖𝑜 𝐵𝑖𝑜𝑚𝑎𝑠𝑠)
2.2.5. Tree parameterisation
In the case of the oil palm, the library already exists, therefore, no parameters needed to be collected in
order to use it in the WaNuLCAS model.
1.7. Model validation
A comparison was conducted between the modelled yield results and field data observed on the trial
plots at the Belaga research station. If the comparison suggests that the results produced by the model
are close to the data obtained from the field, the model will be considered valid.
Wageningen University and Research | Results 15
3. Results
3.1. Pepper allometry and biomass distribution
3.1.1. Leaf biomass
The leaf biomass per individual section was determined using the average number of leaves per section and the average leaf biomass per section (Table 8). Plotting the biomass increment with the increase in
height represented by the sections (Figure 10a) gives us a model allowing to establish the leaf biomass increment (g DM) per increase of 50cm. This relationship is described by the equation: 𝑦 = 2.0868𝑥 + 1.9842.
In figure 10b, we observe that the leaf biomass at 0-50cm is lower than at 50-100. The lower leaf biomass at 0-50 is explained by the fact that they are less leaves present in the section (Table 8). A smaller amount of leaves may be explained by the fact that pepper is propagated generatively using cuttings made from orthotropic branches and the first plagiotropic branch does not appear at ground level but higher. As it is the plagiotropic branches that hold the leaves, this results in a smaller number of leaves, thus biomass. The decrease of leaf biomass starting at 100cm onwards (Figure 10b) is explained by two factors; there are less leaves present per section as we look into higher sections and the leaf size is reduced as we look higher into the sections (Table 8). Indeed, the leaf area per individual leaf decreases as it is located higher on the plant and therefore leads to a lower individual leaf biomass. This trait may be phenotypically related.
Between 200 to 250cm, there is an increase in leaf biomass due to an increase in the number of leaves
present. A greater number of leaves may be due to fact that there is more light available at the top of the
pepper plant, inducing the production of leaves. However, the mean leaf size is smaller. Moreover, only 4
plants measured reached the category of 200-250. Thus, the biomass is not representative to the rest of
the plants measured as they were taller. The influence of the four higher plant on the average leaf biomass
of the lower sections was checked. No significant effect was observed thus, we chose to keep these four
plants in the sample batch.
Table 8, Statistical summary of leaf description
Section Parameter Number of leaves (n)
Biomass leaf (g DM)
Leaf Area (cm²)
Specific Leaf Area (g DM cm-2)
0-50 Mean 274.6 0.3603 7.810 0.0473
Standard Deviation 128.65 0.1032 1.8305 0.0079
Min 100 0.0600 4.187 0.0143
Max 531 0.66 12.109 0.0648
n 30* 150 150 150
50-100 Mean 327.4 0.3521 7.610 0.0463
Standard Deviation 93.52 0.0908 1.7202 0.0069
Min 163 0.1100 4.185 0.0187
Max 529 0.6100 12.207 0.0647
n 30* 150 150 150
100-150 Mean 343.0 0.3342 7.279 0.0459
Standard Deviation 88.55 0.0862 1.599 0.0066
Min 183 0.1600 4.243 0.0252
Max 536 0.5900 12.606 0.0626
n 30* 150 150 150
150-200 Mean 274.3 0.3355 7.103 0.0474
Standard Deviation 109.32 0.0926 1.699 0.0084
Min 81 0.14 3.883 0.0255
Max 508 0.60 12.811 0.0842
n 29* 145 145 145
200-250 Mean 340.8 0.2981 6.622 0.0447
Standard Deviation 93.36 0.10 1.607 0.0099
Min 240 0.09 3.519 0.0177
Max 466 0.63 11.318 0.0644
n 4* 145 145 145
*= number of trees on which the number of leaves per section were recorded
Figure 10, (a)Relationship between cumulated leaf biomass and section,
(b) relationship between non-cumulated leaf biomass and section.
Wageningen University and Research | Results 16
3.1.2. Plagiotropic branches biomass
The linear model used to convert the theoretical plagiotropic branch volume to the real branch volume
(equation 3) uses the following function: 𝑦 = 1.442𝑥 + 0.079 , with an R² value of 0.97 (Figure 11a). The
model used to convert the real plagiotropic branch volume to its biomass in dry matter (equation 4) uses
the following formula: 𝑦 = 0.385𝑥 − 0.028 , with an R² value of 0.97 (Figure 11b). Both regression models
are based on n=735 measurements of plagiotropic branches.
The transformation coefficients were used to determine the biomass (g DM) of the branches measured
non-destructively. The model used to estimate the total plagiotropic biomass based on the number of
leaves present per branch biomass uses the following equation: 𝑦 = 3.784𝑥 + 6.126 and allows to explain
54.2% of the variation.
Based on the cumulative leaf biomass (Figure 10a), we can model the plagiotropic branch biomass to
height increment (Figure 13a) defined by the following equation: 𝑦 = 24.252 + 137.321. There is a low
plagiotropic branch biomass in the first 50cm, it may be explained by the fact that pepper is propagated
generatively using cuttings made from orthotropic branches and the first plagiotropic branch does not
appear at ground level but higher. Between 50cm to 150cm, the plagiotropic branch biomass increases.
This is due to a greater amount of branches per sections. The specific branch biomass per sections was
analysed and showed a decrease in biomass as the branch was in higher sections (Appendix 3). We may
therefore conclude that the increase in biomass is related to the increase in the number of branches present
per section. This trait, like the leaf biomass, may be phenotypically related.
Figure 11, Plagiotropic internodes conversion factors. (a) Relationship between theoretical
volume of the internodes and their real volume, (b) Relationship between the internode
real volume and their biomass (DM).
Figure 12, Relationship between the number of leaves
observed per branch and the branch biomass.
Figure 13, (a)Relationship between cumulated plagiotropic branch biomass and
section, (b) relationship between plagiotropic branch biomass and section.
Wageningen University and Research | Results 17
3.1.3. Orthotropic branches biomass
The linear model used to convert the volume of the orthotropic branches to its biomass (g DM) (Figure 14)
uses the following function: 𝑦 = 0.409𝑥 − 0.099. It is based on the analysis of 186 samples of orthotropic
brannches. The volume used in the regression is based on the conversion of the measured diameters at
each intersection using the conical frustum equation.
The relationship between the orthotropic branch biomass and the height at which it is located is following
a decreasing pattern. Indeed, Figure 15b shows that the orthotropic branch biomass is lower at the upper
part of the plant than at the lower sections. This is due to the growth of the plant, where the orthotropic
branches grow in height but in width over time.
The orthotropic branch biomass per individual section was determined using the average values of biomass
per section (Appendix 4). Plotting the orthotropic branch biomass increment with the increase in height
represented by the sections (Figure 15a) gives us a polynomial model of order 2 allowing to establish the
orthotropic branch biomass increment (g DM) per increase of 50cm. This relationship is described by the
equation: 𝑦 = −0.039𝑥2 + 18.256𝑥 − 225.853.
Figure 14, Orthotropic branch conversion factor.
Relationship between the real volume and the
biomass (DM).
Figure 15, Relationship between cumulated orthotropic branch biomass and section, (b)
relationship between orthotropic branch biomass and section.
Wageningen University and Research | Results 18
3.1.4. Fruit biomass
The best relationship found to predict the fruit biomass per section is through the number of leaves present
per section, with an R2 of 0.62. The linear model (Figure 16) described as 𝑦 = 0.305𝑥 + 11.272 allows us to
predict the number of genererative organs present per section. The number of fruits present per given
section is multiplied by the average fruit biomass (Table 9).
The relationship between the increase in fruit biomass and plant height is constant (Figure 17a). The linear
model is expressed by the following equation: 𝑦 = 1.845𝑥 − 9.408. The trend of the fruit biomass according
to the height follows a bell-shaped pattern, where the fruit biomass per section increases until 150cm high.
The fruit biomass decreases after 150cm (Figure 17b). This is due to the fact that only the plagiotropic
branches hold the generative organs. Indeed, the decrease in fruit biomass after 150cm is related to the
decreasing amount of plagiotropic branches present in the section above 150cm as well as to the
decreasing specific biomass of the plagiotropic branches.
Table 9, Statistical summary of fruit description
Parameter Biomass Fruit (g)
Mean 0.8527
Standard Deviation 0.3927
Min 0.05
Max 2.25
n 618
Figure 16, Relationship between the number of leaves and number of generative
organs observed on the 30 plants measured non-destructively.
Figure 17, (a) Relationship between cumulated fruit biomass and section, (b) relationship
between fruit biomass and section.
Wageningen University and Research | Results 19
3.1.5. Biomass distribution
The biomass distribution is dependent on the height of the pepper plant. Indeed, as the distribution of each
organ is not evenly spread throughout the plant, the size of the plant will determine the biomass
distribution (Figure 18). The detailed description of the biomass distribution is found in appendix 5. The
total biomass is of 1853.88 g, 3883.36g, 5717.84g, 7357.32g and 8801g, for the 50, 100, 150, 200 and
250 sections, respectively. We may observe on Figure 18 that the plagiotropic branch biomass proportion
increases with the height increment. On contrary, the orthotropic branch biomass proportion decreases.
This is due to the fact that the plant first allocates most of the photosynthates to the vertical growth of the
plant, thus through the orthotropic branches.
3.1.6. Identification of farm effect
The sampling carried out to create the allometric equations and the biomass distribution took place on
three different farms, where similar farming practices were conducted.
A first step was conducted to see if there is a difference between farmers in the data that was used to
make the pepper model. The data related to the farms and used for the allometric equations calculus are;
number of leaves per section, number of fruits per sections, orthotropic branch biomass per section. The
P-Value were 3.02e-04, 9.01e-06 and 2.59e-04 for the farm effect on leaves, fruits and orthotropic
branches, respectively. We may therefore reject the null hypothesis, stating that farm has no effect on the
three parameters.
A comparison was carried out to compare the different farms in order to assess farm effect. Figure 19
shows the cumulative biomass of all the organs for a given plant of 250cm high. We can observe that the
farm Muksin has the greatest overall biomass. However, when looking at the relative values, we can
observe that the biomass distribution is similar throughout the three farms and the general model
combining the data of all three farms. We may therefore conclude that there is indeed a farm effect
however, there is no effect on the biomass distribution.
0.580 0.589 0.612 0.640 0.673
0.337 0.273 0.248 0.234 0.228
0.663
0
0.674 0.685 0.673
0.235 0.220 0.187 0.021
0.057 0.064 0.059
Figure 18 Stacked bar plot describing the distribution of biomass of the pepper plant. The bar plot on the left describes the relative
biomass distribution according to the height increment. The bar plot on the right describes the absolute biomass distribution of
the pepper plant. The darkest grey represents the plagiotropic branch biomass, followed by the orthotropic branch biomass, leaf
biomass and the lightest grey represents the biomass allocated to the fruits.
0.060
Figure 19. The bar plot on the left describes the relative biomass distribution according to the height increment per farm and for
the general model. The bar plot on the right describes the absolute biomass distribution of the pepper plant. The darkest grey
represents the plagiotropic branch biomass, followed by the orthotropic branch biomass, leaf biomass and the lightest grey
represents the biomass allocated to the fruits.
Wageningen University and Research | Results 20
3.2. WaNuLCAS modelling results
The WaNuLCAS model was used to assess the effect of intercropping on oil palm and pepper. The modelling
was performed using WaNuLCAS 4.3.1 on the dynamic modelling platform Stella 7.0.3 Research. The
simulations were done under two conditions; 1) monoculture, using the recommended planting pattern;
2) intercrop, using the double-row avenue system. All simulations using the WaNuLCAS model were done
using the same fertilisation regime, climatic conditions, edaphic conditions and managerial activities.
3.2.1. Oil palm in monoculture
The conventional planting pattern of oil palm was elaborated to optimise the canopy area. Indeed, the
conventional planting configuration allowed for a closed canopy, leading to an optimum area allowing the
capture of the photosynthetic active radiations. Figure 20 shows the modelled Fresh Fruit Bunch production
per annum, under the conventional planting pattern. The production of fruit starts 4 to 5 years after
planting. Figure 20 shows that the production curve follows a normal FFB production pattern, where the
immature phase takes place for 4 years after planting, followed by the young phase, where the yield
reaches its peak and finally, followed by a plateauing stage. The average yield at the mature stage is of
20.5 Mg FFB ha-1 year-1. Such values and production patterns are described in literature.
The double-row avenue pattern is not configured to optimize the canopy area. Indeed, the reduction of
space between the palm lines, leads to a smaller ground cover allocated per tree, thus a reduced canopy
area able to use the PAR. Figure 21 shows the fraction of light used by the palm trees. We can observe
that the fraction in the conventional planting pattern is greater in comparison to the double-row avenue
system. The opening in the avenues, in the double-row avenue system, could have had the effect to
compensate this decrease in canopy area. Indeed, the absence of competition for space allows the palm
fronds to capture lateral light, thus leading to a greater area for PAR capture. However, this was not
observed in the modelling results. Figure 20 shows that planting the palms under the double row avenue
system pattern, without any crop integrated into the opening, induces an average yield at the mature
stage of 18Mg ha-1 year-1. We may therefore conclude that the effect of additional light captured in the
openings of the avenue is not sufficient to compensate for the decrease in canopy area.
Figure 20, Oil palm production curves (Mg ha-1 year-1). The circles and plain line represent yield under the conventional
planting pattern. The triangles and the dotted line represent yield under the double-row avenue pattern.
Figure 21, Fraction of light received by the tree. The circles represent the light fraction used under the conventional
planting pattern. The triangles represent the light fraction used under the double-row avenue pattern.
Wageningen University and Research | Results 21
3.2.2. Pepper in monoculture
The simulation of pepper was conducted by integrating the research results in the WaNuLCAS model. The
allometric equations and biomass distribution were used to establish the data set used in the Crop Library
in order to be able to simulate the growth and fruit yield of pepper, under various conditions. The simulation
of pepper in monoculture has only one planting pattern. The planting pattern of pepper is the same for
both in the conventional pattern and when using the double-row avenue system. Moreover, the maximum
height of pepper was set in the model at 250cm, as it is in a pepper field maintained under the best farming
practices. The time span of the simulation for pepper was 25 years, in order to follow the plantation cycle
of oil palm.
Figure 22 describes the production curve of pepper. We can observe that the beginning of production
occurs after three years. After 5 years, the production reaches its plateau. This may be explained by the
fact that the pepper plant grows in height at the beginning, by allocating more of its biomass to the
orthotropic branches. Once the plant has reached its final height, there is an increase in total biomass,
until it has reached its maximum. No production curve has been found in literature. May we therefore not
confirm the production curve established by the model. However, the yield at mature production is on
average at 3.1Mg DM ha-1 year-1. The simulated result using the WaNuLCAS model corresponds with the
yields reported in literature (Ravindran, 2003).
3.2.3. Oil palm and pepper under the double-row avenue system
Intercropping may lead to competition for resources. Figure 23 shows a decrease in yield of FFB when
pepper is added to the plantation. The average yield at mature stage is of 15.5 Mg ha-1 year -1. This may
be due to the competition for nutrients and water. Indeed, both the oil palm and the pepper are sourcing
their nutrients and water from the same niche, leading to competition for nutrients and water. However,
the stress induced by the change in nutrient availability was not expressed by the WaNuLCAS model.
Figure 22, Pepper production over time (Mg ha-1 year-1). The curve represents the
production curve.
Figure 23, Oil palm production curves (Mg ha-1 year-1). The circles and plain line represent oil palm yield under the conventional
planting pattern. The triangles and the dotted line represent oil palm yield under the double-row avenue pattern. The squares
and finely dotted line represent oil palm yield when combined with pepper under the double-row avenue system.
Wageningen University and Research | Results 22
It is assumed that oil palm and pepper compete for nutrients. However, the competition for light is more
pronounced when looking at the shading effect of the palms on pepper. Figure 24 shows, in the initial stage
of the intercrop, that pepper has a yield corresponding to the yield observed when grown in monoculture.
We may qualify the effect of shading of the established palms to be minim on the production of pepper for
5 years after planting. However, the shading effect of the palms increases (Figure 25b) with the increase
in palm trunk height, frond length and crown radius (Figure 25a) has a significant effect on the pepper
production, located in zone 3 (Figure 7). The pepper located in zone 4 (Figure 7) is less affected by the
growth of palm. This may be due to the fact that the pepper plants located in zone 4 are less affected by
the shade. Aravindakshan et al., 1969, report that yield is dependent on the amount of photosynthates
available.
Figure 25a shows that the length of the fronds t greater than the radius of the crown. This is may be due
to the fact that the WaNuLCAS model took into account the fact that the petiole of the fronds does not
attach to the trunk horizontally, thus creates an arc from the bending frond.
Figure 25b describes the amount of light that is used by the pepper plants in zone 3 and 4. We observe
that the amount of light captured by the pepper plant is starting to decrease after 5 years of plantation.
This effect is in concordance to the fact that the palm leaf and thus, the palm canopy radius are increasing
their size. After 5 years after plantation, the palm fronds and radiuses are of 4.98m and 4.57m respectively.
The canopy of the palm starts, after 5 years only, to extend into zone 3 (Figure 7), which is allocated to
pepper. After 19 years after planting, the amount of light captured in zone 3 is 0.2g m-2. Therefore, pepper
plant growth is impossible at this stage.
The amount of light captured by pepper in zone 4 starts to decrease 19 years after planting. This
corresponds to the time when the palm crown radius is 6.43m, thus starts to extend onto zone 4. Syakir,
1994 states that pepper yield pepper is affected when the available light goes below 25% of shading. In
case of the zone 4, the 25% decrease of available light corresponds to 23 years after planting however,
the modelled results show a yield decrease at around 15 years.
Figure 24, Pepper production (Mg ha-1 year-1). The circles and plain line represent yield in monoculture. The squares and the
dotted line represent yield under the double-row avenue pattern, in zone 4. The triangles and finely dotted line represent yield
under the double-row avenue system pattern, in zone3.
Figure 25, (a) Trunk height increase over time (circles), frond length increase (squares) and crown radius (triangles) increase
over time. (b) Amount of light intercepted in zone 3 (circle) and zone 4(triangles)
Wageningen University and Research | Discussion 23
4. Discussion
Using the WaNuLCAS model allowed us to characterize and assess the growth of oil palm and pepper under
the double-row avenue system, for a period of 28 years.
We observed in the modelled results that the production of oil palm decreases by an average of 3.5 Mg ha-
1 year-1 when the planting pattern is changed from conventional to the double-row avenue system. Goh,
1982 reported that there is a decrease in yield of FFB per palm corresponding to the increase in plantation
density. Indeed, adopting the double-row avenue system induces an increased density of 192 palms ha-1
within the row. Moreover, the expected increase in light interception, due to the additional radial light
coming through the openings of the avenues, creating a border effect, did not compensate the decrease
of yield related to the increase in density mentioned by Goh, 1982.
A decrease of 2.5 Mg ha-1 year-1 of oil palm production yield was observed when intercropping oil palm and
pepper. However, we did not witness any stress related to the competition for water or nutrients when
pepper was intercropped with oil palm under the double-row avenue system.
The effect of intercropping oil palm and pepper had a significant effect on the production of pepper. The
pepper row closest to the palm rows was the most affected by the palms. The shade induced by the palm
trees lead to significant decrease in production. Pepper plants located in the middle of the avenue were
less affected by the palm shade. This supports the idea that light had the greatest impact on pepper
production. Moreover, the fact that the pepper plants closer to the palms are more affected in comparison
to the pepper plants located in the middle of the avenue corroborates with the study of Mohd Haniff et al.,
2003 and Suboh et al., 2009, stating that there is more photosynthetically active radiation available for
the plants as we get closer to the centre of the avenue.
As proposed by Nuertey et al., 2000, one advantage of intercropping is the reduction of the financial burden
of replanting oil palms. Indeed, for the farmer, replanting induces a loss of income due to no production
by the palms and a waiting period for the first production of fruit bunches. Despite the high value of pepper,
it may not be the right crop to compensate the financial burden induced by replanting in the short term.
Indeed, pepper takes a few years to attain its mature yield. This means that the farmers may not be able
to rely on pepper only as a source of income. However, once the production of pepper has started, the
financial income is may play an important role in the farmer’s income. Prices are reported to be at around
3000US$ ton-1 for black pepper an around 5000US$ ton-1 for white pepper (International Pepper
Community, 2018).
In the long term, the application of intercropping oil palm and black pepper under the double-row avenue
system involves challenges. Indeed, pepper can be produced in the avenue for around 10 years. However,
the decrease in pepper yield induced by the shade of the palms affects greatly the production of pepper
located closest to the palm rows.
The data obtained through modelling was compared to the data published by Zulkifli et al., 2016, obtained
by a field trial where oil palm and black pepper are intercropped under the double-row avenue system.
The values obtained for oil palm production through modelling are in line with the values obtained by
Zulkifli et al., 2016 however, we may only compare the fresh bunch production for a period of 4 years as
their publication only mentions 4 years of production. Our study provides additional information regarding
the effectiveness of oil palm production in the long-term effect of intercropping. The modelled results
concerning the pepper production corroborate with the results from Zulkifli et al., 2016. Indeed, as their
experiment only lasted for 4 years after the first production of fresh bunches on the palms, they did not
yet reach the time frame needed to see the effect of shading on the pepper plants located closest to the
palms, thus concluded a satisfactory result for the long-term production of pepper. However, our study
leads to refute the conclusion stated by Zulkifli et al., 2016, where black pepper is a suitable crop for
integration in the double-row avenue system as it is not technically feasible through the entire lifespan of
the palm tree.
Various management practices are possible to encounter with the decrease in pepper yield. Farmers may
choose to grow pepper in the entire avenue and replace all pepper plants or the closest one to the palm
rows by a sciaphilic crop after ten years. Another option may be to use oil palm varieties with shorter fronts
however, this may impact the oil palm yield.
Wageningen University and Research | 24
Wageningen University and Research | Conclusion 25
5. Conclusion
This study shows the challenges met when implementing intercropping as it induces a more complex
system. However, it offers to the farmer a large range of choices. In addition to the oil palm production,
the farmer may choose to plant a cash crop, a subsistence crop or both. These managerial strategies offer
the farmer the ability to be less dependent on the market prices and may offer more food security.
Intercropping oil palm also induces a greater diversification within the plantation, which may have
beneficial effects on the surrounding environment. Intercropping in oil palm may be a promising system
allowing to improve the current practices on many aspects.
This study stresses out the importance and advantages of modelling as it allows us to do long term
predictions. The modelling tools are an opportunity to skip years of trials and come to faster conclusions
regarding the best management practices to adopt when intercropping. Therefore, it could allow for a
faster change within the oil palm industry, if the institutes, governments and NGOs follow on closing down
the future knowledge gap between researchers and farmers.
Wageningen University and Research | 26
Wageningen University and Research | Reference 27
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Wageningen University and Research | Appendix 1 29
Appendix 1
Parameters Units Source
Crop Stage
Length of generative stage days Literature
Length of vegetative stage days Literature
Annual or Perennial
Is it annual crop? 1=yes, 0 = no [] Literature
Earliest day to flower in a year Julian day Literature
Latest day to flower in a year Julian day Literature
Crop Growth
Production of dry matter per day kg m-2.day-1 Literature
Seed weight kg m-2 Literature
Water requirement for dry matter production l kg-1 Literature
Ratio of height increment to biomass incr. m kg-2 Literature
Maximum proportion of crop biomass remobilized as storage component
1/day Literature
Light Capture
Extinction light coefficient [] ??
Relative light intensity at which shading starts to affect crop growth [] ??
Canopy
Maximum Leaf Area Index [] Field measurement
Rain Interception
Rainfall water stored at leaf surface mm Field measurement
Water Uptake
Hydraulic conductivity of roots cm day-1 ??
Maximum plant potential cm ??
Minimum plant potential cm ??
Nutrient Uptake
Canopy biomass for closed canopy kg m-2 Field measurement
N concentration in crop tissue [] Field measurement
P concentration in crop tissue [] Field measurement
N concentration in young crop biomass [] Field measurement
P concentration in young crop biomass [] Field measurement
N concentration in roots [] Field measurement
N Fixation N fixation
Type of N2 fixation [] Literature
Constant
Proportion of N from atmosphere [] Literature
Responsive to N stress
Responsiveness of N2 fix. to N stress [] Literature
Fraction of reserve pool for N2 fix. 1/day Literature
Dry weight cost for N2 fixation kg(DW) g(N)-1 Literature
Wageningen University and Research | Appendix 1 30
Parameters Units Source
Root
All root type
Root tip diameter cm Field measurement
Root type 0
Max. root length density in layer1 cm cm-3 Field measurement
Max. root length density in layer2 cm cm-3 Field measurement
Max. root length density in layer3 cm cm-3 Field measurement
Max. root length density in layer4 cm cm-3 Field measurement
Root type 1
Total root length per unit area cm cm-2 Field measurement
Root type 1 and 2
Decrease of root with depth 1/m Field measurement
Root type 2
Length per unit root dry weight m g-1 Literature
Root half life days Literature
Root affected by water or nutrient stress [] Literature
Root distribution by depth in good uptake [] Literature
Mycorrhizal fraction
Fraction of roots infected by mycorrhiza [] Field measurement
Crop effects on nutrient mobility
Reduction of constant P by root activity mg cm-1
Relative transfer rate for N pool cm2 day-1
Relative transfer rate for P pool cm2 day-1
Litter Quality
Lignin fraction of crop residue [] Literature
Lignin fraction of crop root residue [] Literature
Polyphenol fraction of crop residue [] Literature
Polyphenol fraction of crop root residue [] Literature
Soil erosion
Crop cover efficiency factor [] Field measurement
Sensitivity to pest damage
Fraction of crop eaten by pigs [] MPOB
Fraction of crop eaten by monkeys [] MPOB
Fraction of crop eaten by locust [] MPOB
Fraction of crop eaten by nematode [] MPOB
Fraction of crop eaten by goat [] MPOB
Fraction of crop eaten by buffalo [] MPOB
Fraction of crop eaten by birds [] MPOB
Agronomic yield
Standard moisture content [] []
Wageningen University and Research | Appendix 2 31
Appendix 2 Macro for Leaf Area:
run(“Smooth”);
setOption (“BlackBackground”, false);
run (“Make Binary”);
run (Set Scale…”, “distance=294.6 known=1 unit=cm”);
run (“Fill Holes”);
run (“Create Selection”);
run (“Measure”);
rin (“Summarize”);
saveAs (“Results”, xxx);
The macro may be typed directly in the batch process window in ImageJ.
Appendix 3 Statistical summary of the number of leaves recorded and the average biomass and leaf area per section
Section Parameter Plagiotropic Branch Biomass (g DM)
0-50 Mean 41.679
Standard Deviation 26.1817
Min 7.539
Max 165.659
n 145
50-100 Mean 47.016
Standard Deviation 24.7429
Min 8.506
Max 126.348
n 150
100-150 Mean 44.02
Standard Deviation 22.1671
Min 5.00
Max 107.78
n 150
150-200 Mean 36.300
Standard Deviation 17.4943
Min 3.916
Max 84.508
n 145
200-250 Mean 33.166
Standard Deviation 16.4345
Min 8.699
Max 66.875
n 20
*= number of trees on which the number of leaves per section were recorded
The above boxplot shows the decrease in biomass per specific branch per increase in height on the plant.
Wageningen University and Research | Appendix 4 32
Appendix 4
Statistical summary of the orthotropic branch biomass per section
Section Parameter Orthotropic Branch Biomass (g DM)
0-50 Mean 589.7
Standard Deviation 304.9927
Min 129.9
Max 1328.3
n 30
50-100 Mean 612.9
Standard Deviation 276.52
Min 173.7
Max 1230.1
n 30
100-150 Mean 450.23
Standard Deviation 233.2053
Min 32.65
Max 932.72
n 30
150-200 Mean 193.48
Standard Deviation 195.4213
Min 1.55
Max 723.89
n 30
200-250 Mean 58.24
Standard Deviation 66.6770
Min 2.63
Max 215.04
n 30
Appendix 5
Pepper biomass distribution per height (g)
Height Total Biomass Plagio Biomass Ortho Biomass Leaf Biomass Fruit Biomass
50 1853.88 1075.28 589.45 106.32 82.83
100 3883.36 2287.88 1209.75 210.66 175.07
150 5717.84 3500.48 1635.05 315.00 267.31
200 7357.32 4713.08 1865.35 419.34 359.55
250 8801.80 5925.68 1900.65 523.68 451.79