Anais da Academia Brasileira de Ciências (2018) 90(4): 3265-3283

(Annals of the Brazilian Academy of Sciences)

Printed version ISSN 0001-3765 / Online version ISSN 1678-2690
http://dx.doi.org/10.1590/0001-3765201820160806
www.scielo.br/aabc | www.fb.com/aabcjournal

Plant growth, radiation use efficiency and yield of sugarcane cultivated

in agroforestry systems: An alternative for threatened ecosystems

FELIPE SCHWERZ1, SANDRO L.P. MEDEIROS2, ELVIS F. ELLI3, ELDER

ELOY4, JAQUELINE SGARBOSSA5 and BRAULIO O. CARON5

1
Departamento de Produção Vegetal, Escola Superior de Agricultura Luiz de Queiroz/ESALQ,
Universidade de São Paulo/USP, Avenida Páduas Dias, 11, 13418-900 Piracicaba, SP, Brazil
2
Departamento de Produção Vegetal, Universidade Federal de Santa Maria,
Avenida Roraima, 1000, 97105-900 Santa Maria, RS, Brazil
3
Departamento de Engenharia de Biossistemas, Escola Superior de Agricultura Luiz de Queiroz/ESALQ,
Universidade de São Paulo/USP, Avenida Páduas Dias, 11, 13418-900 Piracicaba, SP, Brazil
4
Departamento de Engenharia Florestal, Universidade Federal de Santa Maria, Campus de Frederico
Westphalen, Linha Sete de Setembro, s/n, BR 386, Km 40, 97105-900 Frederico Westphalen, RS, Brazil
5
Departamento de Ciências Agronômicas e Ambientais, Universidade Federal de Santa Maria, Campus de Frederico
Westphalen, Linha Sete de Setembro, s/n, BR 386, Km 40, 97105-900 Frederico Westphalen, RS, Brazil

Manuscript received on November 25, 2016; accepted for publication on November 6, 2017

ABSTRACT
Sugarcane (Sacharum officinarum L.) monocropping has had a great socio-economic and environmental
impact in Brazil, and agroforestry systems have been considered as an alternative for more sustainable
production; however, there is a lack of field research under such conditions. The aim of this study was to
evaluate the growth rates, radiation use efficiency and yield traits in sugarcane cultivated in the understory
of Aleurites fordii, in two agroforestry arrangements and monocropping systems. A field experiment was
conducted from July 2015 to June 2016 in the city of Frederico Westphalen, Rio Grande do Sul, Brazil.
The radiation use efficiency, assimilate partitioning, leaf area index, absolute growth rate, net assimilation
rate, number of tillers, plant height, % of intercepted solar radiation, extinction coefficient, and yield
in each system was evaluated. In agroforestry systems, the dynamic interactions between multiple plant
species change with the time and can result in unique microclimates. The use of agroforestry systems in 12
x 12m arrangements should be prioritized because it enables greater yields and radiation availability in the
understory. This study sought to provide new sustainable alternatives for farmers in order to increase the
diversification of the rural property and maintain the preservation of existing agroecosystems.
Key words: biomass partitioning, crop systems, growth rates, Sacharum officinarum, yield traits.

INTRODUCTION and environmental preservation. In order to achieve

this balance, it is necessary to meet the demand for
One of the greatest national and global challenges food and energy without compromising existing
is to generate a balance between crop production agroecosystems (Godfray et al. 2010). Agroforestry
Correspondence to: Felipe Schwerz systems deserve to be highlighted in this scenario
E-mail: [email protected] and are a promising strategy in order to achieve these

An Acad Bras Cienc (2018) 90 (4)

3266 FELIPE SCHWERZ et al.

objectives. These systems consist of integrated land such as solar radiation (Muller et al. 2014, Elli et
uses, for example, land used for forestry purposes, al. 2016). The amount of solar radiation transmitted
crop production, and/or raising livestock. Such through the canopy can be presented in direct or
systems provide clear agro-ecological advantages diffuse forms, which is a determining factor in the
over systems in which only one crop is grown internal microclimate of a system (Pezzopane et
(Brooker et al. 2014). Advantages include higher al. 2015). Direct and diffuse radiation can affect
production per unit of land (Zhang et al. 2007, Li et radiation use efficiency and light extinction
al. 2013), greater resource-use efficiencies of water coefficients (Campbell and Norman 1998), which in
and nutrients (Vandermeer 1989, 2011), greater
turn may modify the growth rate (Pinto et al. 2005),
carbon sequestration in the soil (Makumba et al.
assimilation and partitioning of photoassimilates
2006, Cong et al. 2014), and an increased input of
produced by species in the understory (Mendes et
organic matter, which has been shown to improve
al. 2013). In the case of sugarcane, the response due
chemical, physical and biological properties of soil
to reduced radiation available within the canopy
(Tracy and Zhang 2008, Salton et al. 2013).
can be optimized. Sugarcane, a C4 metabolism
Intercropping is a dominant strategy of
plant, has a high solar radiation demand, in order to
agriculture in many parts of the world, i.e. sub-
meet the photosynthetic rates of plants.
Saharan Africa and large parts of Latin America,
Biomass production in plants depends upon
and provides an estimated 20% of the world’s food
the quantity of absorbed photosynthetically active
supply (Altieri 2009, Chappell et al. 2013). Despite
radiation (PARa) by leaves, and the efficiency by
the great potential of intercropping, little research
which leaves convert and assimilate the radiant
has been carried out to determine the traits of
light through photosynthesis. Thus, the intercepted
cultivated species which drive the positive effects
photosynthetically active radiation (PARi) that is
of intercropping systems.
converted into biomass reveals the efficiency of the
Brazil is one of the major sugarcane producing
use of radiation (εb) by a species (Monteith 1977,
countries in the world, with estimates for 2015/2016 Van Heerden et al. 2010). While photosynthesis is the
of 8.97 million hectares in planted area, which yields basis for the production of biomass, little attention
685 million tons of sugarcane, with an average has been given to the radiation use efficiency in
yield of 76 tons ha-1 (Conab 2017). Sugarcane order to improve sugarcane yields (Zhu et al. 2010,
(Sacharum officinarum L.) monocropping has Flood et al. 2011). Solar radiation intercepted by
had a great socio-economic and environmental the plant canopy is the most important component
impact in Brazil, and agroforestry systems (AFs) for growth analysis, but to estimate this radiation,
have been considered as an alternative for the it is necessary to know the LAI and the extinction
sustainable production in threatened ecosystems. coefficient.
Despite the potential for AFs previously described, Solar radiation can change at different points in
little research has been carried out to assess plant the understory of agroforestry systems (Paciullo et
growth, yield traits, and the dynamics of resources al. 2011), depending on the area of shade provided
available in these systems. by the canopy, and the orientation and spacing of
The dynamic interactions between plants in the trees. Thus, new projects should be developed
agroforestry systems are especially relevant in areas in order to evaluate the growth, yield, and radiation
in which plants grow beneath trees, given that a use efficiency of sugarcane cultivated in different
tree’s height, crown projection, and leaf area index agroforestry arrangements, as well as in different
can influence the distribution of existing resources positions within each system. There is a lack of

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3267

published results for field experiments under such evaluation of sugarcane occurred in September
conditions. 2015 and the last in May 2016. The sugarcane
To address this lack of information, the utilized was developed by the Agronomic Institute
following hypothesis was created: The cultivation of Campinas (IAC), cultivar IAC 87-3396, which is
of sugarcane in agroforestry systems provides characterized by a high yield, sucrose content, and
greater yield, radiation use efficiency, and plant excellent adaptation to soils with lower fertility.
growth, and thus can be recommended to farmers as In the 12 x 12 system, forest species were
a sustainable alternative of production in threatened distributed in lines spaced at 12m. The sugarcane was
ecosystems. The aim of this study was to evaluate distributed in eight lines arranged in correspondence
the growth rates, radiation use efficiency, and yield to each interval between the lines of forest species,
traits of sugarcane cultivated in the Aleurites fordii totaling 16 lines throughout the system. In the 6 x
understory, in two agroforestry arrangements and a 6 system, the forest species was planted in lines
monocropping system. spaced at 6m. The sugarcane was distributed in
four lines in each interval corresponding with lines
MATERIALS AND METHODS of Aleurites fordii plants. A total of 15 trees were
STUDY AREA AND EXPERIMENTAL DESIGN allocated to each experimental unit. The forest
species were planted in the experimental field in
A field experiment was conducted from July 2015 to September, while the sugarcane was planted in
June 2016 in the city of Frederico Westphalen, Rio November 2011. After plowing and harrowing
Grande do Sul, Brazil, at the coordinates 27º23’48’’ seedlings were manually planted. In both systems,
S, 53º25’45’’ W and an altitude of 490 m. According the sugarcane had a spacing of 1.20m and a density
to Köppen’s climate classification (Alvares et al. of 16 buds per meter, with both tree and sugarcane
2013), the climate is Cfa, i.e., humid subtropical lines oriented from East to West. After planting
with a mean annual temperature of 19.1°C, and the sugarcane, plots were delineated by stakes set
varying maximum and minimum temperatures of two meters apart and were distributed at different
38ºC and 0°C, respectively. The experiment was points in the understory of each experimental unit.
performed in ratoon sugarcane (1st cut performed In the 12 x 12 system lines L1, L2 and L3 of the
on 07/26/2012), and the analyses were conducted evaluation were positioned at a distance of 1.2m,
in the 4th cut, crop cycle 2015/2016. The soil of 3.6m and 4.8m from the Aleurites fordii species,
the experimental area was classified as typical respectively. Differently, in the 6 x 6 system lines
Oxisol. Fertilization was carried out in response L1, L2 and L3 were positioned at 1.2m, 2.4m and
to a soil analysis following the recommendations 1.2m, respectively. These plot areas were chosen
for sugarcane crops by soil chemistry and fertility with the objective of representing microclimate
committee (SCFC 2004). conditions in areas under the canopy of each
The experimental design was a randomized agroforestry system. For subsequent analyses of the
complete block, characterized by a factorial data, average values of the lines in each system were
arrangement of 3 x 3 x 8, with three cropping calculated in order to comply with the objectives of
systems, two agroforestry systems (12 x 12 and this study. The arrangement of tree, sugarcane, and
6 x 6) and an isolated system with sugarcane; plot of evaluation are shown in Figure 1.
three positions in line of the agroforestry systems Forest species Tung (Aleurites fordii), which
(Line 1, Line 2 and Line 3); and eight months of belongs to the Euphorbiaceae family, an exotic
plant collection, with three repetitions. The first species with deciduous characteristics, was used

An Acad Bras Cienc (2018) 90 (4)

3268 FELIPE SCHWERZ et al.

Figure 1 - A sketch of an experimental unit of the agroforestry systems (12 x 12 and 6 x 6)

and the isolated system. Black circles represent the trees, continuous lines indicate where the
sugarcane was planted, and the rectangles in grey represent the monthly evaluation plots of
sugarcane.

to establish the agroforestry system. It was chosen In the laboratory, plant sectioning was
because of its adaptability to environmental performed, as well as the preparation of leaf discs to
conditions and high oil content in its fruits, which determine the leaf area and dry matter partitioning.
make it a viable option for the composition of The total dry matter (TDM) of the plants was
agroforestry systems. Allometric characteristics determined as the sum of the components: leaf,
of Aleurites fordii trees were evaluated using pseudoculm+senescent, and stalk. Each component
descriptive statistics (Table I). was gathered and placed in labeled, individual
paper sacks. The sacks were then sent into a forced
PLANT GROWTH EVALUATIONS
circulation oven at 60°C until a consistent mass
The sugarcane was cut on July 2, 2015; evaluations was obtained. The samples were later weighed on
started 84 days after cutting (DAC), due to the a precision balance in order to obtain the dry mass
slow initial growth of shoots, evaluations were of each component, which together resulted in the
performed every 30 days, with the last at 332 TDM.
DAC, amounting to 8 months of plant collection. Leaf area (LA) was calculated according to the
In each evaluation period, a total of 42 plants equation:
were collected, and two representative plants per ( DM leaves+discs )
LA = ( nº discs*punch disc area ) *
evaluation plot were analyzed, resulting in a total DM discs
of 332 evaluated plants. On the day of collection,
tiller numbers were counted and plant height was Where, n° discs = number of discs by sample;
measured with a metric tape measure. punch area = punch disc area in mm2; DM leaves

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3269

TABLE I
Height (H), stem diameter (SD), diameter at breast height (DBH) and mean diameter of crown (MDC) of Aleurites fordii,
in agroforestry systems (12 x 12 and 6 x 6), the plants were five years old.

H (m) SD (cm) DBH (cm) MDC (m)

Statistics
12 x 12 6x6 12 x 12 6x6 12 x 12 6x6 12 x 12 6x6

Average 3.46 3.78 20.02 26.19 11.58 14.50 1.79 2.60

Standard
91.02 113.14 6.08 6.69 3.71 3.32 74.11 144.73
Deviation
Variance 1285.50 1801.14 36.96 44.73 13.74 11.03 1492.30 2946.49
Kurtosis 3.03 -1.70 -2.72 -0.46 -0.71 -0.76 0.70 1.74
Minimum 269.00 225.00 12.60 17.60 6.80 10.20 91.50 113.00
Maximum 500.00 524.00 26.20 36.50 16.40 19.60 289.50 536.50

= total dry matter of the leaves, in grams; and DM The extinction coefficient (k) was calculated
discs = dry matter of the disc, in grams. The leaf area using the following equation:
index (LAI) from the total leaf area of each plant and
Rn
the soil area (SA) occupied by plants was determined ln ( )
k=- Rt
according to the equation: LAI = LA/SA. LAI

Based on the results of dry matter and leaf area, Where k = extinction coefficient; Rn = solar
the growth analysis of the variables was conducted: radiation measured under the plant canopy (MJ
total dry matter (TDM, g m-2), absolute growth rate m-2); Rt = radiation above the plant canopy (MJ
(AGR, g day-1); net assimilation rate (NAR, g m-2 m-2); LAI = leaf area index.
day-1) according to the methodology described in The values for intercepted global radiation
the literature (Thornley 1976, Marafon et al. 2012). (IGR) were measured monthly, where the incident
radiation was measured above and under the
EXTINCTION COEFFICIENT AND LIGHT
plant canopy with a portable pyranometer, which
INTERCEPTION
recorded measurements in the period from 10 to
Solar radiation at each evaluation period was 12h. The values of intercepted global radiation
measured using a portable sensor pyranometer were obtained according to the following equation:
(LICOR PY32164) coupled to a Datalogger
% Intercepted = [100 - (Rn x 100 / Rt)]
(LICOR 1400). The values of incident global solar
radiation were obtained from the meteorological Where: Rn = incident radiation under the
station of the National Meteorology Institute, canopy; Rt = incident radiation above the canopy.
situated roughly 200m from the experiment. The
RADIATION USE EFFICIENCY
values of active photosynthetically radiation were
estimated to be 45% of global solar radiation. Production of dry matter was based on the
This fraction follows the average values found model proposed by Monteith (1977), whereby
by Assis and Mendez (1989). The estimation of the dry matter production was calculated from
accumulated photosynthetically active radiation intercepted photosynthetically active radiation
was based on the methods by Monteith (1977) and (PARi) multiplied by the use efficiency. The εb
Varlet-Grancher et al. (1989). was calculated by the relation between the average

An Acad Bras Cienc (2018) 90 (4)

3270 FELIPE SCHWERZ et al.

production of accumulated TDM and the PARi differences were found at 5% probability among
involved in the production of biomass according to the cropping systems. We reject the null hypothesis
the following expression: H0. The Dunnet test (p> 0.05) was used to compare
the 12 x 12 and 6 x 6 intercropping systems with
TDM = εb * PARi
the control (monocropping system) and the Tukey
Where TDM = total dry matter produced (g test (p> 0.05) to compare the difference between
-2
m ); PARi = intercepted photosynthetically active intercropping systems 12 x 12 and 6 x 6. In
solar radiation (MJ.m-2); and εb = radiation use addition, a regression analysis for the variable of
efficiency of dry matter produced (g. MJ-1). The intercepted global radiation was performed.
value of the radiation use efficiency given by
RESULTS
the angular coefficient represents the amount of
accumulated biomass for each unit of intercepted METEOROLOGICAL CONDITIONS
energy.
Values for intercepted photosynthetically The air temperature during the sugarcane crop
active radiation were based on the model proposed cycle ranged from 6.3ºC to 31.0ºC, with an average
by Varlet-Grancher et al. (1989): temperature of 19.2ºC; the flux of global solar
radiation was 15.89 MJ.m-2.dia-1 on average, with a
PARi = 0.95 * (PARinc) * (1- e (-k * LAI))
variation of 1.98 to 33.03 MJ.m-2.dia-1; the rainfall
Where: PARi = intercepted photosynthetically accumulated during the crop cycle was 2587.5mm
active radiation (MJ.m -2); PARinc = incident (Figure 2).
irradiant photosynthetically active radiation (MJ.m- RADIATION USE EFFICIENCY AND LIGHT
2
); k = extinction coefficient, which was set to INTERCEPTION
0.20 for all systems, as this was the average value
obtained in this study; LAI = leaf area index. Growth of TDM presented a positive linear
relationship with PARia during the crop cycle and
YIELD TRAITS AND STATISTICAL ANALYSIS presented high correlation coefficients (Figure
3a). We observed variations in the radiation use
The yield traits were evaluated at 350 DAP, by
collecting and weighing the sugarcane stalk efficiency of the conversion into biomass, in
samples from each plot, and each of the twelve different cropping systems and evaluation lines
different positions in each system, in order to within the agroforestry systems.
obtain greater homogeneity of the samples. The Of the crop systems, the greatest radiation
yield was determined on the basis of the variables use efficiency of sugarcane was obtained by the
stalk weight and juice volume. The stalk weight isolated system (2.28 g MJ-1). In relation to the
(SW, t ha-1) was obtained with the aid of a digital evaluation lines, we observed that for the 12 x 12
scale. The juice volume (JV, m3 ha-1), was obtained system, line 3 showed a greater efficiency, 2.03 g
from milling the stalk and measuring the obtained MJ-1, when compared to the lines L1 and L2, which
juice with the aid of a graduated cylinder with a showed values of 1.89 and 1.85 g MJ-1, respectively,
capacity of 1 L. revealing a 9% difference for plants located in the
The data were statistically analyzed with center of the plot. For the 6 x 6 system, we observed
the software “Statistical Analysis System” a greater radiation use efficiency in line 2, which
(SAS 2003). According to the F test, significant demonstrated 1.96 g MJ-1, i.e., 10 and 6% higher

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3271

Figure 2 - Monthly average values of minimum, maximum and average temperature, accumulated
incident solar radiation, and accumulated rainfall during the experimental period (07/02/2015 to
06/24/2016).

than the values in line 1 (1.77 g MJ-1) and line 3 (1.84 LIGHT EXTINCTION COEFFICIENT
g MJ-1) of the evaluation, respectively (Figure 3a).
The k values were similar in the different cropping
The intercepted global radiation showed
systems (Table II). In the initial stages of plant
a quadratic response for the different cropping
growth, k values were 0.15 on average. Due to
systems and evaluation lines (Figure 3b). The
the plant growth, through the elevated LAI and
greatest amount of intercepted global radiation
dry matter accumulation (Figure 4), the extinction
was observed for the isolated system of sugarcane,
coefficient showed increased values, with a
which intercepted 89.3% of incident radiation at
maximum value of 0.32 at 237 DAC. Considering
237 DAC. In the agroforestry systems, we found
the crop cycle and generalizing the crop systems,
different response values for each evaluation line.
the average light extinction coefficient for the
In the 12 x 12 system, we observed a reduction of sugarcane crop was 0.20.
16.5 and 12.5% in the interception of radiation with
the sugarcane lines nearest to the forest species (L1 LEAF AREA INDEX AND DRY MATTER
PRODUCTION
and L2), when compared to L3, which intercepted
85.4% of the radiation at 265 DAC. Furthermore, The highest values of LAI were observed in the
the 6 x 6 system showed a similar response, where stage of stalk growth, a period between 237-
lines L1 and L3 presented lower values of 13 and 265 DAC. The isolated system had the highest
8.5% in the interception of radiation compared LAI, 7.1; additionally, the intercropping system
to the L2 evaluation line, which showed 82.3% provided changes in values of LAI in accordance
interception of the incident solar radiation at 265 with the line position. The 12 x 12 system showed
DAC (Figure 3b). the highest value in line 3, with a value of 6.2,

An Acad Bras Cienc (2018) 90 (4)

3272 FELIPE SCHWERZ et al.

TABLE II response curves demonstrated similar behavior for

Light extinction coefficient with (± standard error) in
the analyzed variables. We were able to observe two
different cropping systems of sugarcane cultivation
throughout the days after cutting (DAC). peaks during the crop cycle; the first occurred at 200
DAC and the second at 300 DAC, with a subsequent
Cropping Systems
DAC decrease in growth rates in accordance with the
12 x 12 6x6 Monocrop senescence of the leaves and stalk maturation.
84 0.142 ± 0.038 0.124 ± 0.066 0.115 ± 0.047 In addition, a similar response in the different
115 0.158 ± 0.031 0.135 ± 0.092 0.136 ± 0.056 crop systems and lines of evaluation was verified.
145 0.163 ± 0.039 0.169 ± 0.021 0.145 ± 0.037 Overall, the largest absolute growth rate and net
174 0.190 ± 0.016 0.190 ± 0.029 0.159 ± 0.020 assimilation rates were observed at 206 DAC, this
206 0.211 ± 0.038 0.275 ± 0.071 0.261 ± 0.019
stage corresponds with intense stalk growth.
237 0.302 ± 0.024 0.324 ± 0.066 0.337 ± 0.018
The number of tillers in sugarcane reduces in
265 0.285 ± 0.067 0.296 ± 0.044 0.295 ± 0.026
accordance to the DAC, stabilizing at 265 DAC
(Figure 6a). Overall, a similar response between
300 0.207 ± 0.011 0.217 ± 0.033 0.228 ± 0.049
the crop systems and lines of evaluation throughout
332 0.190 ± 0.026 0.145 ± 0.031 0.183 ± 0.023
the crop cycle were observed. In the initial growth
stages, a period of intense tillering, we observed
which is 15.6 and 12.5% higher than in lines 1 and
an average of 10 tillers m -2. In the stage of
2, respectively. In the 6 x 6 system, the highest
maturation, we found values between 4 to 5 tillers
LAI was found in line 2, with values around 6.4;
m-2, a reduction of more than 50% in the number
these values were 14.5 and 11.6% higher than those
of tillers. In relation to plant height (Figure 6b),
obtained by lines 1 and 3, respectively (Figure 4a).
we observed a linear response in accordance to the
Dry matter accumulated during the cycle of
sugarcane presented a positive linear relation, DAC, with stabilization in height from 300 DAC
with an observed stabilization for 300 DAC. This onwards. Overall, the plants presented values of
period marks the beginning of the maturation of approximately 265 cm.
stalks, which can be noted by the reductions in BIOMASS PARTITIONING
growth rates and an increase in sucrose content
of stalks (Figure 4b). The greatest values for total The pattern of dry matter accumulation in the leaves,
accumulated dry matter were observed in the pseudoculm+senescent, and stalks of sugarcane
isolated system, which increased a total of 592.7 was similar for the different cropping systems
g plant-1. For the intercropping systems, a similar (Figure 7). In the initial stages of plant growth, up
response in accumulated dry matter was obtained to 115 DAC, the leaves and pseudoculm+senescent
from the different evaluation positions in the were responsible for 100% of total dry matter
understory. While the 12 x 12 system had a total of accumulation, 50% for each compartment on
504.8 g plant-1 on average, the accumulation total average; the dry matter of the stalks was computed
of dry matter of the 6 x 6 system was found to be from 115 DAC onwards. At 206 DAC the plants
469.3 g plant-1 (Figure 4b). showed stabilization in the partition of assimilates
up to the harvest; where the leaves (20%)
GROWTH RATES
pseudoculm+senescent (5%) and stalk (75%) were
The absolute growth rate and net assimilation rate responsible for the total dry matter accumulated by
throughout the crop cycle are shown in Figure 5. The the sugarcane crop.

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3273

Figure 3 - The relationship between accumulated intercepted photosynthetically active radiation

(PARiac) and dry matter produced (DM) (a); and intercepted global radiation of sugarcane in
agroforestry and isolated systems in different lines of evaluation throughout the days after cutting (b).

YIELD TRAITS higher than those observed in the agroforestry

systems (Figure 8). We observed a reduction of
According to the variance analysis, we observed a 17.5 and 32.7% of stalk weight for the 12 x 12
significant difference for the variables stalk weight and 6 x 6 systems, respectively, when compared
and juice volume in the different cropping systems to the isolated system by the Dunnet test (p> 0.05).
(Figure 8). The stalk weight and juice volume Additionally, between the crop systems, the stalk
in sugarcane grown in the isolated system were yield was 18.4% greater in the 12 x 12 system

An Acad Bras Cienc (2018) 90 (4)

3274 FELIPE SCHWERZ et al.

Figure 4 - Leaf area index (a) and total dry matter (b) of sugarcane in agroforestry and isolated systems in different lines
of evaluation throughout the days after cutting.

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3275

than in the 6 x 6 according to the Tukey test (p> In addition, excessive shading during extended
0.05). In relation to the juice volume produced, a periods may affect the photosynthetic apparatus as
similar response to the stalk weight was observed, a result of decreases in chlorophyll content (Chl),
confirming the correlation between these variables, in nitrogen content, and in the Chl a/b ratio (Pearcy
where the isolated system also showed a higher 1998, Dinç et al. 2012).
juice volume. We also observed a reduction of 16.6
The values of radiation use efficiency obtained
and 35.3% by juice volume produced in the 12 x
in this study are similar to what has been observed
12 and 6 x 6 systems, respectively. In addition,
by other authors: De Silva and De Costa (2012)
between agroforestry systems, the juice volume of
the 12 x 12 system was 22.5% higher than in the 6 found values of RUE which vary from 1.63 to 2.09
x 6 system. With this, we can infer that the 12 x 12 g MJ-1 and 0.71 to 1.03 g MJ-1 under irrigated and
system showed greater stalk yields and volume of rain-fed conditions in Sri Lanka; Anderson et al.
produced juice, compared to the 6 x 6 system. (2015), analyzing the RUE in two crop sites in
Hawaii, USA, found average values of 1.24 ± 0.22
DISCUSSION g MJ-1 during the period of growth for locations of
low altitude and 1.15 ± 0 15 g MJ-1 for location of
The meteorological conditions during the growing
high altitude. The maximum RUE for sugarcane
cycle were favorable for sugarcane, especially with
respect to the air temperature and water availability. can vary between 1.7 g MJ-1 (Robertson et al. 1996)
Sugarcane needs 1500 to 2500mm of water evenly to 2.0 g MJ-1 (Muchow et al. 1997).
distributed over the growing season (FAO 2013). In the national and international literature,
Within the agroforestry system the positioning RUE values for sugarcane grown in agroforestry
of sugarcane lines is very important, and therefore systems have not been determined. Comparing
should be considered in studies with intercropping the RUE values obtained from the monocropping
systems; the canopy of forest species can result system with those obtained in the intercropping
in changes to microclimates. In this case, the systems in this study, even under the influence of
canopy provided a reduction in LAI (Figure the tree species, the sugarcane in the understory
4a) and consequently in intercepted radiation of agroforestry system showed high values for
(Figure 3b) in the lines of sugarcane near the tree RUE and can, therefore, be recommended for
species, resulting in an overall lower radiation use cultivation in alternative production systems, such
efficiency by plants. The growth and development as agroforestry systems.
of different species in the same area cause dynamic The lowest amount of radiation intercepted by
interactions in species planted near one another, the sugarcane in the 12 x 12 system, in the first and
which changes with the time. These interactions second lines of evaluation, as well as in the first and
modify the distribution of existing resources in the third lines in the 6 x 6 system, are associated with
system (Muller et al. 2014), with the solar radiation the effects of the canopy of the species Aleurites
changing first. fordii, which shaded the sugarcane plants. Due to
The results observed are consistent with those sugarcane’s high photosynthetic rate (C4 species),
of Marchiori et al. 2014, who reported that excessive it needs a large amount of solar radiation to meet
shading may affect certain growth traits causing its photosynthetic demand. Plants submitted
decreases in tillering and in photosynthetically to different levels of shading (for example, in
active leaf area, as well as in the interception of proximity of the evaluation line to the tree species)
solar radiation, producing thinner, elongated stalks. demonstrate changes in photosynthetic rate, and

An Acad Bras Cienc (2018) 90 (4)

3276 FELIPE SCHWERZ et al.

Figure 5 - Absolute growth rate (a) and net assimilation rate (b) of sugarcane in agroforestry and isolated systems in
different lines of evaluation throughout the days after cutting.

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3277

Figure 6 - Number of tillers (a) and plant height (b) of sugarcane in agroforestry and isolated
systems in different lines of evaluation throughout the days after cutting.

consequently, the production of assimilates, which of the existing gaps in the canopy, which can
influences the radiation use efficiency of plants. all be affected modified by plant arrangement.
The observed results are similar to those of Naturally, the probability of photosynthetically
Hardy et al. (2004), who reported that the amount active radiation reaching the ground is inversely
of radiation intercepted by the forestry species and proportional to the degree in which it is intercepted
consequently the radiation that reaches the ground by the canopy structure.
level is determined by canopy characteristics, Péllico Netto et al. (2015), evaluating the
such as the average crown diameter and the size dynamics of solar radiation in the understory of

An Acad Bras Cienc (2018) 90 (4)

3278 FELIPE SCHWERZ et al.

Acacia mearnsii, concluded that in the center of the Inman-Bamber et al. (2011) in a study conducted in
plot, i.e. between rows of trees, more than 70% of the Australia with a value of 0.58. It can be highlighted
radiation reaches the ground level. However, under that in both national and international literature, k
the canopy, the values were reduced significantly, values were not determined for sugarcane grown in
to only 20%, thereby diminishing the availability agroforestry systems.
of radiation for understory crops. This confirmed The greatest values of LAI were found in
the influence of the canopy trees on the interception periods of intense vegetative growth (237 to 265
of incident radiation. Our findings are in line with DAC), with values near to 7. In accordance with the
a study carried out by Pezzopane et al. (2015) and results of Inman-Bamber (1994), for the cultivation
Bosi et al. (2014) who reported the strong relation of sugarcane, the critical value of LAI to intercept
between levels of incident solar radiation, and its 95% of the radiation is near 4, thus average values
effect on microclimate, growth characteristics of obtained in this study are above the critical value
plants, and soil moisture. of LAI.
The growth of sugarcane was correlated to an According to Pinto et al. (2005), the main
increase in k values (Table II), coinciding with an limiting factor in agroforestry systems is the
increase in LAI. This result provides an increase availability of solar radiation, which together with
in radiation interception up to a determined value the competition for water and nutrients limits the
when the plants reach a critical IAF and begin growth of sugarcane near the forest species. In new
self-shading, resulting in an increased extinction research conducted with agroforestry systems, the
coefficient. It can be inferred that independent of effect of a reduction in radiation availability must
the cropping system, the k values depend primarily be considered due to the influence in the growth
on the LAI, which determines the amount of solar and development of the species in intercropping
radiation available in the understory of sugarcane. systems.
According to Behling et al. (2015), this condition is The changes in the microclimatic conditions
understandable since light attenuation of the plant in different cropping environments resulted in
canopy is determined by leaf density (which can variations of the absolute growth rate and net
be expressed by LAI) and also by the geometric assimilation rate of plants (Figure 5). The reduction
characteristics of the leaves and the tree canopy, as in growth rates is explained by the effect of self-
well as the optical properties of the leaves. Thornley shading due to an increase in the LAI (Figure 4a), as
(1976) reported that the value of k may vary with well as in the ripening stage due to leaf senescence
leaf traits, sun angle, spacing, and latitude. as a result of reduced air temperature. In accordance
High values of LAI (> 6) resulted in the average with the results of Cardozo and Sentelhas (2013),
k value of 0.20, for the conditions as observed in in the south of Brazil, the lowest air temperature in
this study. Similar values were obtained by De the months of autumn-winter, combined with the
Silva and De Costa (2012), who found values occurrence of moderate water deficit, are the main
near to 0.27 in irrigated sugarcane in Sri Lanka, factors responsible for the reduction of growth rates,
although other authors found values higher than except, at the beginning stage of maturation where
those obtained in this study. Muchow et al. (1994, it results in a high increase in sucrose content.
1997) reported k values near to 0.40 for irrigated The reduction in the number of tillers up to
sugarcane in Australia and Hawaii. Meki et al. DAC 237 is due to the effect of intraspecific
(2015) found a value of k equal to 0.53 for sugarcane competition among plants, especially in the initial
in Hawaii. This is similar to a value obtained by stages of growth, i.e., the stage of high tillering. The

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3279

competition occurs mainly for solar radiation since of biomass throughout the sugarcane cycle, found
this is the limiting factor in the agroforestry system. higher initial biomass accumulated in the leaves
In addition, we recorded low levels of cumulative and in stalks starting from 129 DAP. This is
rainfall in August and September, periods that equivalent to 9% of TDM with increasing values,
corresponded to approximately 50 and 80 DAC reaching 70% at 330 DAP. In a study conducted in
and the tiller emission stage, which affected the southern Brazil, Simões et al. (2005) found ratios
tillering of sugarcane. of around 80% of dry matter accumulated in stalks
The tillering pattern found in this study is at 400 DAP. Muchow et al. (1994) found the stalk
similar to that described by Bezuidenhout et al. biomass values change from 60% to 80% for DAP
(2003), but with a lower tiller density than what 300 to 450 DAP, in Australia.
was reported in that study: peak tillering at 12 to 14 The greatest stalk yield and juice volume
tiller m-2. After the senescence stage, tiller density observed for the monocropping system is explained
stabilized at 7 tiller m-2 regardless of water source by the favorable meteorological conditions during
(rain or irrigation) or planting site. This implies the crop cycle (Figure 2). The average stalk yield
that tillering rate and survival of tillers is related of 61.8 t ha-1 is consistent with the values found
to the quantity and quality of the solar radiation in the literature. In the study conducted by Liu et
intercepted by plants (Inman-Bamber 1994, al. (2016), who evaluated the yield of sugarcane in
Bezuidenhout et al. 2003). China, observed an average yield of 56.1 and 79.9 t
Regarding the partitioning of assimilates, ha-1 for Kaiyuan and Yuanjiang, respectively. Abreu
the biomass accumulated in the leaves decreased et al. (2013), studying five cultivars of sugarcane,
gradually throughout the crop cycle, especially in found the following average yields: 89 t ha-1 in the
the maturation stage due to the senescence of lower first year of cultivation, 75 t ha-1 in the second year
leaves. Another important aspect to be emphasized and 88 t ha-1 in the third year of cultivation in the
is the exportation of sucrose and starch from leaves state of Alagoas, Brazil.
to other compartments. In our study, the total dry The sugarcane yield is reduced in agroforestry
matter in the stalk in the sugarcane plants resulted systems. This reduction in stalk weight and juice
in an average accumulation of 75% (Figure 7). volume may be related to the morphophysiological
The metabolism and partitioning of sucrose adjustments, for example as a shade tolerance
are essential for all stages of the sugarcane crop strategy. These adaptations were not able to
cycle and requires hydrolysis to use it as a source compensate for the reduction of radiation in the
of energy (Leite et al. 2009). In leaves (the source), understory of Aleurites fordii, which reduced overall
the synthesis of carbohydrates is realized, which yields. However, even under the influence of tree
is translocated to the stalks (sink) in the form of species, the 12 x 12 system showed satisfactory
sucrose, in order to meet plant maintenance and values with relation to the stalk weight (50.9 t ha-
1
growth metabolism demands (Taiz and Zeiger ) and produced juice volume (26.8 m3 h-1). This
2013). These processes are accompanied by value is near the average stalk yield for the state of
constant changes in source-sink relations (Roitsch Rio Grande do Sul of 53.9 t ha-1 (Conab 2017). This
and González 2004), which was confirmed in this result confirms the viability of the intercropping
study through the partitioning of dry matter by the system for the production of sugarcane. Pinto et
sugarcane (Figure 7). al. (2005), evaluating the yield of sugarcane in
The observed results are similar to those of agroforestry systems with Eucalyptus grandis,
Marin et al. (2011), who, evaluating the partition found differences in sugarcane yields in different

An Acad Bras Cienc (2018) 90 (4)

3280 FELIPE SCHWERZ et al.

Figure 7 - Biomass partitioning in sugarcane plants in different cropping systems

throughout the days after cutting. Each bar represents the mean value of twelve replications
(± s.e.).

evaluation spacings. Plants spaced 11.6 m from the can increase the radiation intercepted by sugarcane,
tree species obtained average yields of 64.1 t ha-1 in especially in the final stage of maturation, resulting
the state of São Paulo, Brazil. in an increase of the stalk weight and produced juice
The satisfactory yield values in the 12 x 12 (Figure 8). This result highlights the importance
system are related to the greater availability of of appropriate selection of tree species used in the
radiation within the system, which enabled a composition of the agroforestry system.
greater plant growth, confirmed by the elevated These results are consistent with those obtained
TDM and LAI (Figure 4). In addition, the Aleurites by Elli et al. (2016), who, evaluating the yield of
fordii (deciduous species), displays leaf senescence sugarcane in different agroforestry systems, found
in the colder seasons (winter-autumn). Senescence higher stalk yields for intercropping with the

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3281

also confirms the viability of the cultivation of

sugarcane in agroforestry systems. Additionally,
Brazilian law No. 12,651, May 25, 2012, which
established a new Forest Code, allows small
farmers to plant agroforestry systems in areas of
permanent preservation (APP) and legal reserves
(RL), provided that those systems are subject to a
sustainable management plan.

CONCLUSIONS

The cultivation of sugarcane in agroforestry system

did not present higher yields and radiation use
efficiency, thus, the hypothesis of the study was
rejected. However, even under the influence of the
species Aleurites fordii, the cultivation of sugarcane
in agroforestry systems showed satisfactory
values of yield and radiation use efficiency and
can be recommended for cultivation in alternative
production systems. The use of agroforestry
systems with 12 x 12m arrangements should be
prioritized because this enabled greater yields and
radiation availability in the understory.
Figure 8 - Stalk weight and juice volume in sugarcane This study sought to provide new sustainable
plants in different agroforestry and monocropping systems.
alternatives for farmers, in order to increase the
Each bar represents the mean value of twelve replications (±
s.e.). Means followed by the same letter do not differ among diversification of rural properties and preserve
themselves. Uppercase compare the agroforestry systems with existing agro-ecosystems. It is important that
the control (Monocropping), by Dunnet test (p> 0.05); and research is conducted in order to study the
lowercase compare the agroforestry systems (12 x 12 and 6 x
6m), by Tukey test (p> 0.05). implementation of agroforestry systems in
threatened ecosystems, such as in permanent
species Parapiptadenia rígida, where the values preservation areas (APP’s) and legal reserve (RL’s).
of 61.9, 53.1, and 41.6 t ha-1 were obtained in three It is necessary to study the responses of these areas
cultivation cycles. The greater yields observed in with regard to environmental preservation, the
by the authors is related to the characteristics of maintenance of biodiversity, and economic return
their forest species. Above all, because they enable of agroforestry systems, and to do so while adapting
greater radiation availability in the understory, an to modern environmental legislation.
important characteristic of deciduous species.
ACKNOWLEDGMENTS
The study of the growth variables, yield traits,
and dynamics of solar radiation in agroforestry The authors wish to acknowledge the Conselho
systems are underfunded. Thus, the information Nacional de Desenvolvimento Científico e
generated in this study is relevant, as it provides Tecnológico (CNPq-Brazil) and the Coordenação
information to farmers for the planning of de Aperfeiçoamento de Pessoal de Nível Superior
more effective agroforestry arrangements. It (CAPES-Brazil) for their financial support.

An Acad Bras Cienc (2018) 90 (4)

3282 FELIPE SCHWERZ et al.

REFERENCES CONG WF, HOFFLAND E, LI L, SIX J, SUN JH, BAO

XG, ZHANG FS AND VAN DER WERF W. 2014.
ABREU ML, ALMEIDA SM, TEODORO I, HOLANDA LA Intercropping enhances soil carbon and nitrogen. Glob
AND NETO GDS. 2013. Crescimento e produtividade de Chang Biol 21: 1715-1726.
cana-de-açúcar em função da disponibilidade hídrica dos DE SILVA ALC AND DE COSTA WAJM. 2012. Growth and
Tabuleiros Costeiros de Alagoas. Bragantia 72(3): 262- radiation use efficiency of sugarcane under irrigated and
270. rain-fed conditions in Sri Lanka. Sugar Tech 14(3): 247-
ALTIERI MA. 2009. Agroecology, small farms, and food 254.
sovereignty. Monthly Review 61(3): 102-113. DINÇ E, CEPPI MG, TÓTH SZ, BOTTKA S AND
ALVARES CA, STAPE JL, SENTELHAS PC, MORAES G, SCHANSKER G. 2012. The chl a fluorescenceintensity
LEONARDO J AND SPAROVEK G. 2013. Köppen’s is remarkably insensitive to changes in the chlorophyll
climate classification map for Brazil. Meteorol Z 22: 711- content of the leaf as long as the chl a/b ratio remains
728. unaffected. Biochim Biophys Acta 1817(5): 770-779.
ANDERSON RG, TIRADO-CORBALÁ R, WANG D AND ELLI EF, CARON BO, ELOY E, BEHLING A, SOUZA VQ
AYARS JE. 2015. Long-rotation sugarcane in Hawaii AND SCHWERZ F. 2016. Productive, morphological and
sustains high carbon accumulation and radiation use qualitative characteristics of sugarcane in the understory
efficiency in 2nd year of growth. Agric Ecosyst Environ tree species in agroforestry systems. Afr J Agric Res
199: 216-224. 11(17): 1576-1584.
ASSIS FN AND MENDEZ MEG. 1989. Relação entre FAO - FOOD AND AGRICULTURAL ORGANIZATION.
radiação fotossinteticamente ativa e radiação global. Pesq 2013. Water, Natural Resources Management and
Agropec Bras 24(7): 797-800. Environment Department. Available at: http://www.fao.
BEHLING A, SANQUETTA DR, DALLA CORTE AP, org/nr/water/index.html. Accessed on June 29, 2016.
NETTO SP, RODRIGUES AL, CARON BO AND SIMON FLOOD PJ, HARBINSON J AND AARTS MGM. 2011.
AA. 2016. Tracking leaf area index and coefficient of light Natural genetic variation in plant photosynthesis. Trends
extinction over the harvesting cycle of black wattle. J For Plant Sci 16(6): 327-335.
Res 27(6): 1211-1217. GODFRAY HCJ ET AL. 2010. Food security: the challenge
BEZUIDENHOUT CN, O’LEARY GJ, SINGELS A AND of feeding 9 billion people. Science 327(5967): 812-818.
BAJIC VB. 2003. A process-based model to simulate HARDY JP, MELLOH R, KOENIG G, MARKS D,
changes in tiller density and light interception of sugarcane WINSTRAL A, POMEROY JW AND LINK T. 2004.
crops. Agric Syst 76(2): 589-599. Solar radiation transmission through conifer canopies. Agr
BOSI C, PEZZOPANE JRM, SENTELHAS PC, SANTOS Forest Meteorol 126: 257-270.
PM AND NICODEMO MLF. 2014. Produtividade e INMAN-BAMBER NG. 1994. Temperature and seasonal
características biométricas do capim-braquiária em sistema effects on canopy development and light interception of
silvipastoril. Pesq Agropec Bras 49(6): 449-456. sugarcane. Field Crop Res 36(1): 41-51.
BROOKER RW ET AL. 2014. Improving intercropping: a INMAN-BAMBER NG, JACKSON PA AND HEWITT M.
synthesis of research in agronomy, plant physiology and 2011. Sucrose accumulation in sugarcane stalks does not
ecology. New Phytol 206: 107-117. limit photosynthesis and biomass production. Crop Pasture
CAMPBELL GS AND NORMAN JM. 1998. The light Sci 62: 848.
environment of plant canopies. In: An Introduction to LEITE GHP, CRUSCIOL CAC, LIMA GPP AND SILVA
Environmental Biophysics. Springer New York, p. 247- MDA. 2009. Reguladores vegetais e atividade de invertases
278. em cana-de-açúcar em meio de safra. Cienc Rural 45(10):
CARDOZO NP AND SENTELHAS PC. 2013. Climatic 718-725.
effects on sugarcane ripening under the influence of
LI L, ZHANG L AND ZHANG F. 2013. Crop mixtures and
cultivars and crop age. Sci Agric 70(6): 449-456.
the mechanisms of overyielding. In: Encyclopedia of
CHAPPELL MJ ET AL. 2013. Food sovereignty: an alternative
paradigma for poverty reduction and biodiversity Biodiversity. Waltham, MA, USA: Academic Press, p.
conservation in Latin America. F1000 Research 2: 235- 382-395.
253. LIU J ET AL. 2016. Growth and yield of sugarcane genotypes
C O N A B - C O M PA N H I A N A C I O N A L D E are strongly correlated across irrigated and rainfed
ABASTECIMENTO. 2017. Acompanhamento da environments. Field Crop Res 196: 418-425.
Safra Brasileira de Cana-De-Açúcar. Disponível MAKUMBA W, JANSSEN B, OENEMA O, AKINNIFESI
em: http://www.conab.gov.br/OlalaCMS/uploads/ FK, MWETA D AND KWESIGA F. 2006. The long-
arquivos/16_08_18_12_03_30_boletim_cana_ term effects of a gliricidia–maize intercropping system in
portugues_-_2o_lev_-_16-17.pdf. Acesso em 14 de junho southern Malawi, on gliricidia and maize yields, and soil
de 2016. properties. Agric Ecosyst Environ 116: 85-92.

An Acad Bras Cienc (2018) 90 (4)

 AN APPROACH ABOUT SUGARCANE IN AGROFOREST SYSTEMS 3283

MARAFON AC. 2012. Análise quantitativa de crescimento em tropical region of Brazil. Agric Ecosyst Environ 105(1):
cana-de-açúcar: uma introdução ao procedimento prático. 77-86.
Documentos - Embrapa Tabuleiros Costeiros, p. 29. ROBERTSON MJ, WOOD AW AND MUCHOW RC. 1996.
MARCHIORI PE, MACHADO EC AND RIBEIRO RV. 2014. Growth of sugarcane under high input conditions in
Photosynthetic limitations imposed by self-shading in tropical Australia. Radiation use, biomass accumulation
field-grown sugarcane varieties. Field Crop Res 155: 30- and partitioning. Field Crop Res 48(1): 11-25.
37. ROITSCH T AND GONZÁLEZ MC. 2004. Function and
MARIN FR, JONES JW, ROYCE F, SUGUITANI C, regulation of plant invertases: sweet sensations. Trends
DONZELI JL AND NASSIF DS. 2011. Parameterization Plant Sci 9(12): 606-613.
and evaluation of predictions of DSSAT/CANEGRO for SALTON JC, MERCANTE FM, TOMAZI M, ZANATTA JA,
Brazilian sugarcane. Agron J 103(2): 304-315. CONCENÇO G, SILVA WM AND RETOREA M. 2013.
MEKI MN ET AL. 2015. Two-year growth cycle sugarcane Integrated crop livestock system in tropical Brazil: toward
crop parameter attributes and their application in modeling. a sustainable production system. Agric Ecosyst Environ
Agron J 107(4): 1310-1320. 190: 70-79.
MENDES MMS. 2013. Desenvolvimento do milho SAS. 2003. Getting started with the SAS Learning Edition.
sob influência de árvores de pau branco em sistema Cary, 200 p.
agrossilvipastoril. Pesq Agropec Bras 48(10): 1342-1350. SCFC - SOIL CHEMISTRY AND FERTILITY COMMITTEE.
MONTEITH JL. 1977. Climate and the efficiency of crop 2004. Manual de adubação e calagem para os estados
production en Britain. Philos T Roy Soc B 181: 277-294. do Rio Grande do Sul e Santa Catarina. Porto Alegre:
MUCHOW RC, EVENSEN CI, OSGOOD RV AND Sociedade Brasileira de Ciência do Solo – Núcleo Regional
ROBERTSON MJ. 1997. Yield accumulation in irrigated Sul, p. 400.
sugarcane: Utilization of intercepted radiation. Agron J SIMÕES MS, ROCHA JV AND LAMPARELLI RAC. 2005.
89(4): 646-652. Growth indices and productivity in sugarcane. Sci Agric
MUCHOW RC, SPILLMAN MF, WOOD AW AND
62(1): 23-30.
THOMAS MR. 1994. Radiation interception and biomass
TAIZ L AND ZEIGER E. 2013. Plant Physiology. 5th ed., Porto
accumulation in a sugarcane crop grown under irrigated
tropical conditions. Aust J Agric Res 45(1): 37-50. Alegre: Artmed, p. 918.
MULLER MD, PACIULLO DSC, MARTINS CE, ROCHA THORNLEY JMH. 1976. Mathematical models in plant
WSD AND CASTRO CRT. 2014. Desenvolvimento physiology. Academic Press, London, p. 318.
vegetativo de pinhão manso em diferentes arranjos de TRACY BF AND ZHANG Y. 2008. Soil compaction, corn
plantio em sistemas agrossilvipastoris. Pesq Agropec Bras yield response, and soil nutrient pool dynamics within
49(7): 506-514. an integrated crop-livestock system in Illinois. Crop Sci
PACIULLO DSC, GOMIDE CAM, CASTRO CD, 48(3): 1211-1218.
FERNANDES PB, MÜLLER MD, PIRES MFA, VANDERMEER JH. 1989. The Ecology of Intercropping.
FERNANDES EM AND XAVIER DF. 2011. Cambridge, UK: Cambridge University Press, p. 241.
Características produtivas e nutricionais do pasto em VANDERMEER JH. 2011. The Ecology of Agroecosystems.
sistema agrossilvipastoril, conforme a distância das Boston, MA, USA: Jones and Bartlett Publishers, p. 387.
árvores. Pesq Agropec Bras 46(10): 1173-1186. VAN HEERDEN PDR, DONALDSON RA, WATT DA AND
PEARCY RW. 1998. Acclimation to sun and shade. In: SINGELS A. 2010. Biomass accumulation in sugarcane:
Raghavendra AS (Ed), Photosynthesis: A Comprehensive
unravelling the factors underpinning reduced growth
Treatise. Cambridge University Press, Cambridge, p. 250-
phenomena. J Exp Bot 61(11): 2877-2887.
263.
VARLET-GRANCHER CG, GOSSE M, CHARTIER H,
PÉLLICO NETTO S, SANQUETTA CR, CARON BO,
SINOQUET R, BONHOMME R AND ALLIRAND JM.
BEHLING A, SIMON AA, CORTE APD AND
1989. Mise au point: rayonnement solaire absorbé ou
BAMBERG R. 2015. Ground level photosynthetically
active radiation dynamics in stands of Acacia mearnsii De intercepté par un couvert végétal. Agronomie 9(5): 419-
Wild. An Acad Bras Cienc 87: 1833-1845. 439.
PEZZOPANE JRM, BOSI C, NICODEMO MLF, SANTOS ZHANG L, VAN DER WERF W, ZHANG S, LI B AND
PM, CRUZ PGD AND PARMEJIANI RS. 2015. SPIERTZ JHJ. 2007. Growth, yield and quality of wheat
Microclimate and soil moisture in a silvopastoral system and cotton in relay strip intercropping systems. Field Crop
in southeastern Brazil. Bragantia 74(1): 110-119. Res 103: 178-188.
PINTO LFG, BERNARDES MS, STAPE JL AND PEREIRA ZHU XG, LONG SP AND ORT DR. 2010. Improving
AR. 2005. Growth, yield and system performance photosynthetic efficiency for greateryield. Annu Rev Plant
simulation of a sugarcane–eucalyptus interface in a sub- Biol 61: 235-261.

An Acad Bras Cienc (2018) 90 (4)