Efeito de Diluição de N em Palma

ARTICLE IN PRESS
Journal of
Arid
Environments
Journal of Arid Environments 67 (2006) 357–372
www.elsevier.com/locate/jnlabr/yjare
Productivity of Opuntia ficus-indica (L.) Miller

under different N and P fertilization and plant
population in north-east Brazil
J.C.B. Dubeux Jr.a,, M.V. Ferreira dos Santosa,
M. de Andrade Lirab, D. Cordeiro dos Santosb,
I. Fariasb, L.E. Limab, R.L.C. Ferreiraa
a
Depto. de Zootecnia, Universidade Federal Rural de Pernambuco, Av. Dom Manoel de Medeiros, S/N,
Dois Irmãos, 52171-900 Recife, PE, Brazil
b
Empresa Pernambucana de Pesquisa Agropecuária (IPA), Av. Gal. San Martin, Bonji,
50761-000 Recife, PE, Brazil
Received 26 September 2005; received in revised form 23 February 2006; accepted 23 February 2006
Available online 17 April 2006
Abstract
The arid and semi-arid region occupies 89.3% of north-east Brazil, and livestock farming is one of
the most important activities in this region. Experiments were conducted in four locations in the
semi-arid of Pernambuco, Brazil. The objective was to study N (0, 75, 150, 225, and
300 kg N ha1 yr1) and P fertilizations (0 and 33 kg P ha1 yr1) at two populations (5000 and
40 000 plants ha1) of Opuntia ficus-indica (L.) Mill. Increasing population from 5000 to
40 000 plants ha1 increased productivity in all four sites. Phosphorus fertilization increased dry
matter yield (DMY) only when soil P was o10 mg kg1. Nitrogen fertilization increased linearly
DMY in the higher population, but no response to N was observed with 5000 plants ha1. Nitrogen
fertilization also increased plant N concentration and rainfall use efficiency. Cladode area index
(CAI) increased with population but even in the higher population the CAI was 1 or less. In
summary, increasing plant population and fertilizing with N and P resulted in greater productivity of
Corresponding author. Tel.: +55 81 33206574.

E-mail address: dubeux@ufrpe.br (J.C.B. Dubeux Jr.).
0140-1963/$ - see front matter r 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.jaridenv.2006.02.015
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358 J.C.B. Dubeux Jr. et al. / Journal of Arid Environments 67 (2006) 357–372
O. ficus-indica in north-east Brazil. Phosphorus fertilization involved less economical risk to the
farmer but positive responses occurred only when soil P was o10 mg kg1.
r 2006 Elsevier Ltd. All rights reserved.
Keywords: Cactaceae; CAM; Semi-arid; Spacing; Spineless cactus
1. Introduction
The arid and semi-arid region occupies 89.3% of north-east Brazil (Pereira, 1992), with
livestock production being one important activity in the rural areas. The seasonal forage
production, concentrated within 3–4 months of the rainy season, is a major obstacle to
increase animal productivity in this region. The spineless cactus forage is an important
alternative for farmers because it has demonstrated high productivity potential in this
harsh environment. Despite its low protein concentration, the cactus forage provides
available energy, water, and minerals (Véras et al., 2002). If associated to silage and
concentrate, animals are able to reach higher performance (Melo et al., 2003) as opposed
as when fed alone.
Araújo Filho (1977) suggested that approximately 60% of north-east Brazil should use
xerophitic crops. In fact, spineless cactus is already grown in north-east Brazil and the two
most cultivated species are Opuntia ficus-indica and Nopalea cochenillifera occupying an
estimated area of 400 000 ha in this region (Farias et al., 1984). Farmers have been using
this forage since the 1930s in north-east Brazil (Duque, 1980), mainly in the states of
Pernambuco, Paraı́ba, and Alagoas. A field survey showed that 84.6% of the farmers in
Serra Talhada County, semi-arid region of Pernambuco state, grow spineless cactus in
their properties. In São Bento do Una, ‘Agreste’ region of the same state, the spineless
cactus occupies 32% of the properties area and is considered essential for the dairy activity
(Chagas, 1992).
Because of its CO2 fixation mechanism (CAM), spineless cactus has high water use
efficiency (Viana, 1969). Recent published data showed annual productivity up to
20 t DM ha1 yr1 in Pernambuco semi-arid region (Santos et al., 2000). Considering that
this forage has 750 g kg1 of total digestible nutrients (TDN) (Santos et al., 1990a), it is
difficult to obtain another forage option with the same level of DM productivity and
energy concentration in the semi-arid environment. As any other cut-and-carry system,
however, the soil nutrient extraction from the spineless cactus crop is high. Considering the
average concentration (DM basis) of N, P, K, and Ca as 9, 1.6, 25.8, and 23.5 g kg1,
respectively (Santos et al., 1990b), and the productivity obtained by Santos et al. (2000),
i.e. 20 t DM ha1 yr1 of DM, the annual nutrient extraction per ha by the crop would be
180 kg of N, 32 kg of P, 516 kg of K, and 470 kg of Ca, without considering the other
macro and micronutrients. Therefore, without a fertilization program spineless cactus
productivity will decline over time because of reduction in soil fertility. Nevertheless, an
important aspect of mineral nutrition of this species is not known yet, particularly in the
conditions prevailing in the semi-arid region of north-east Brazil. Increasing productivity
of the spineless cactus will improve the adequacy of this crop for the small farm holder,
which is the prevalent type of farmer in north-east Brazil. The increase in spineless cactus
productivity may also free up some areas in the small property, which may in turn be
cultivated with cash crops like maize, cassava, and beans.
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J.C.B. Dubeux Jr. et al. / Journal of Arid Environments 67 (2006) 357–372 359
The low soil fertility levels and the lack of use of fertilizers contribute to reduce the
forage harvested from cactus field in this region. Organic fertilization is an option and has
been used by some farmers, however, the need of the organic fertilizer for cash crops like
maize and beans reduce the amount available for use in the cactus plantations. Other
important factor reducing the output of forage from this crop is the low plant density
commonly found among the producer’s fields. Thus, the objective of this research was to
evaluate the potential use of N–P fertilization as a viable practice for O. ficus-indica cv.
‘Clone 20’. In order to accomplish that, different levels of N and P were applied in two
populations, 5000 plants ha1 and 40 000 plants ha1. The study was repeated in 4 sites,
with 4 replications per site. Plant DM productivity, plant N concentration, cladode area
index (CAI), rainfall use efficiency (RUE) and, total cladode number were determined. We
hypothesized that N–P fertilization increases DM yield and N concentration of O. ficus-
indica cv. ‘Clone 20’ but interactions with site and plant population are likely to occur.
2. Material and methods
The experiments were carried out at 4 different sites (Arcoverde, São Bento do Una,
Sertânia, and Serra Talhada) located in the semi-arid region of Pernambuco State, north-
east Brazil to evaluate the ‘clone 20’ (O. ficus-indica, Mill.) response to different N–P
fertilization and plant population. Clone 20 is a genetic material obtained by selecting the
free pollination progeny originated from O. ficus-indica cv. ‘Gigante’ (Santos, 1992). The
soil in Arcoverde and São Bento do Una is classified as Regosol (or Entisol). In Sertânia
and Serra Talhada, the soil is classified as luvisol (or Alfisol). The altitude of the four sites
is 429, 558, 614, and 663 m above sea level for Serra Talhada, Sertânia, São Bento do Una,
and Arcoverde, respectively.
Treatments included different sites, N and P fertilization levels, and plant populations. A
split–split plot arrangement was used with the locations (Arcoverde, São Bento do Una,
Sertânia, and Serra Talhada) considered as the main plot, the complete factorial
arrangement among N levels (0, 75, 150, 225, and 300 kg ha1 yr1 N) and P levels (0 and
33 kg ha1 yr1 of P) as the split plot, and the plant populations (5000 and
40 000 plants ha1) as the split–split plot. A randomized complete block design was used
with 4 replications per treatment. The planting spacing was 2 m 1 m (5000 plants ha1)
and 1 m 0.25 m (40 000 plants ha1). The subplot area was 40 m2 (8 m 5 m) and 20 m2
(8 m 2.5 m) with utile area of 12 and 6 m2, for the 5000 and 40 000 plants ha1,
respectively. One border row was excluded on all sides in the 5000 plants ha1 population
and two border rows were excluded in the more dense population. All the cladodes in the
subplots were harvested and weighted.
The O. ficus-indica Mill. cv. ‘Clone 20’ was used and one mature cladode was planted per
pit. All the phosphorus for the 2-year period was applied at the planting date and after the
harvest, using triple super phosphate as P source. The entire N was applied in the first year
of growth, divided in two applications. Potassium was applied with the N, according to the
soil fertility lab recommendations, being the same for all plots. The K source was KCl and
it was applied at a rate of 220 kg KCl ha1 2-yr1. Urea was the N source whereas KCl was
the potassium source of fertilizer. Weed mechanical control (hoe) was performed. At the
harvest (2 yr after planting), the primary cladodes were preserved on all the plants.
Planting and harvesting dates of the different locations are in Table 1. Fresh weight was
recorded and three cladodes representing different cladode orders from each subplot were
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Table 1
Planting and harvesting date of the experiments in the four different sites
Site Planting date Harvest date
São Bento do Una May 12, 1998 April 2000

Arcoverde May 3, 1998 May 2000
Sertânia January 20, 1999 January 18, 2001
Serra Talhada January 4, 1999 November 1, 2000
taken to determine DM concentration (65 1C for 72 h). Cladodes were cut in a longitudinal
section to allow rapid moisture loss. Nitrogen concentration in the cladode was determined
using the Kjeldahl method (Silva, 1981).
CAI was determined according to the equation proposed by Santos (1992) where the
total perimeter of the cladodes correlates with the cladode’s area (Area-
184.96+7.66perimeter; r2 ¼ 0:90; po0:05). The perimeter is the length around the
cladode border and it was taken using a tape measure. The CAI is obtained dividing the
cladode’s area by the area occupied by each plant. Six plants per plot were measured and
the average result was used for the calculations. Evaluations were performed in the
following dates and sites: October 16, 1999 (São Bento do Una), September 28, 1999 (Serra
Talhada) and September 30, 1999 (Sertânia). Total cladode number per plant was
evaluated from 28 to 30 September 1999 in Arcoverde, Sertânia, and Serra Talhada. The
RUE was estimated dividing the harvested DM by the cumulative rainfall during the total
growth period. The cumulative rainfall was 805.8, 1056.9, 965.0, and 1204.4 mm for São
Bento do Una, Arcoverde, Sertânia, and Serra Talhada, respectively. The primary
cladodes were not included in this calculation because they were not harvested.
Statistical analyses were performed using Proc Mixed from SAS Inst. Inc. (1996) and
means compared with the LSMEANS statement. When N fertilization was significant
(p40:05), polynomial orthogonal contrasts (linear, quadratic, and cubic) were performed.
3. Results and discussion
3.1. Dry matter yield
The dry matter yield (DMY) ranged from 3.1 up to 47.3 t DM ha1 harvest1, with the
lowest value observed in Serra Talhada (5000 pl. ha1, 300 kg N ha1 yr1 and 0 kg P) and
the greatest value observed in Sertânia (40 000 pl. ha1, 300 kg N ha1 yr1 and
33 kg P ha1 yr1), respectively. DMY was affected by an interaction between site and
plant population (Table 2). Higher plant population resulted in greater yield in all sites.
Interaction occurred because the increase in yield with increased plant population was
proportionally greater in Serra Talhada than in the other sites (Table 2). Increase in plant
population has been reported in the literature as an effective way to improve Opuntia spp.
productivity (Flores Valdez and Aguirre Rivera, 1992; Medeiros et al., 1997). Greater light
interception, leading to less weeds and higher photosynthetic efficiency per unit land area is
the likely reason for DM yield increase in more dense populations (Dubeux et al., 2000).
The DM yield varied among sites with average values ranging from 6 to 17 t ha1 of DM in
the 5000 plants ha1 density and 19.4–33.7 t ha1 of DM in the 40 000 plants ha1 density.
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Table 2
Dry matter production of ‘Clone 20’ (Opuntia ficus-indica Mill.) at different sites and plant populations in north-
east Brazil
Site Plant population P levela
5000 plants ha1 40 000 plants ha1
t DM ha1 harvest1
Arcoverde 13.8 abb 24.3 b o0.0001

S.B. do Una 9.3 bc 17.8 c o0.0001
Serra Talhada 6.0 c 19.4 bc o0.0001
Sertânia 17.0 a 33.7 a o0.0001
S.E. ¼ 2.3 t DM ha1 harvest1.

a
P level for comparisons within the same row.
b
Means followed by the same letter within the same column are not different (p40:05) by the LSMEANS test.
Considering the environmental conditions that prevailed in these 4 sites, this level of
productivity may be considered appropriate, particularly because of the high digestibility
of this forage (Véras et al., 2005). Because the primary cladodes were not cut at the harvest
time, the primary productivity that occurred during the 2-yr growth was greater than the
numbers presented here.
Plant population also interacted with N fertilization affecting DM yield of Clone 20. No
response (p40:10) to N fertilizer occurred when plant population was 5000 plants ha1 but
a linear response occurred (po0:001) when the plant population was 40 000 plants ha1.
Because of the higher productivity in the higher plant population, N uptake per area by the
crop is also higher, leading to a greater N response for DM yield. Nobel et al. (1987)
concluded that N, P, and possibly B fertilization can increase the growth of Opuntia
engelmannii salm-Dyck and O. rastrera Weber. Gonzalez (1989) also observed a significant
positive effect of N and P fertilization on the DM yield of Opuntia lindheimeri Engelm in a
long-term experiment; however, the author did not find fertilization effect in the first 2 yr
after planting. According to Nobel (1995), the five soil elements that apparently have a
major effect on Opuntia growth are N, P, K, B, and Na. Claassens and Wessels (1997)
found optimum yields with an N application between 30 and 60 kg N, which coincides with
a leaf N concentration of 0.96%. Lima et al. (1974) tested up to 100 kg ha1 2 yr1 of N
applied in O. ficus-indica and observed a positive linear response in the DM yield with a
population of 20 000 plants ha1. Contrasting with these results, Galizzi et al. (2004)
evaluating fruit yield of O. ficus-indica observed significant positive effect for exchangeable
soil Ca, but not for applications of N, P, and K. These authors also highlighted the greater
Ca and K concentration compared to N concentration in the cladodes.
There was an interaction of plant population and P fertilization which also affected DM
yield. Interaction occurred because there was a significant (po0:05) increase in the DM
yield with P fertilization only in the higher plant population, which increased from 21.2 to
26.4 t ha1 harvest1 of DM when P was not applied or at 33 kg ha1 yr1 of P,
respectively. The same explanation for the N fertilization by plant population interaction
hold here, i.e. greater P demand in higher population likely increased Opuntia response to
applied P. Lima et al. (1974) found a linear response of O. ficus-indica up to 22 kg ha1 yr1
of P, stabilizing the yield after that level. Nobel (1995) reported that the P levels that reach
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50% of the maximum yield for Opuntia spp. are relatively low. The same author concluded
that P is not a major limiting factor for Opuntia spp. development. Therefore, a substantial
Opuntia growth may occur in soils with low P levels; however, P mineralization for the
cattle fed with Opuntia is required.
There was a site by P fertilization interaction affecting DM yield (Table 3). The greater
response to P fertilizer occurred in Sertânia followed by Serra Talhada. In the last site the
difference was marginally significative (po0:06). No significant response (p40:10) to P
occurred in the other two sites. When P was added, the greatest DMY occurred in
Sertânia, followed by Arcoverde with the lowest yields occurring in S.B. do Una and Serra
Talhada. The initial soil P levels in Sertânia and Serra Talhada were 9 and 7 mg kg1, while
the initial soil P levels in Arcoverde and São Bento do Una were 28 and 24 mg kg1,
respectively. This might explain the increase in DM yield with P fertilization only in
Sertânia and Serra Talhada. Cavalcanti (1998) recommended 44 kg P ha1 at the planting
date when soil P level iso11 mg dm3 and half of this amount when soil P is in the
11–30 mg dm3 range, for O. ficus-indica with higher plant population (40 000 plants ha1).
According to the same author, no P fertilizer is necessary when soil P level is
430 mg dm3.
Although economic analysis is not the main objective of this research, some
considerations are indeed necessary in terms of technology adoption by small farmers.
In north-east Brazil, spineless cactus is sold during the drought period to feed the cattle
and the prevailing price is US$ 0.09 kg1 of DM. Lira et al. (2005) estimated the
production cost of spineless cactus in north-east Brazil in US$ 0.09 kg1 of DM (i.e. the
same selling price), including in their calculation the inputs such as fertilizer, herbicide, and
labor with cultivation and harvesting. Fertilization cost in Brazil is approximately US$
0.87 kg1 N and US$ 1.00 kg1 P (1 US dollar ¼ 2.29 Brazilian reais). Because there was a
positive linear effect of N only in the higher plant population, our brief economic
considerations are restricted to that plant population. Considering that each kg of added N
produced an average of 15.75 kg DM of extra cactus, that cactus price is US$ 0.09 kg1
DM, and that N cost is US$ 0.87 kg1 N, the N fertilizer represented 61% of the selling
price for the additional forage harvested. This is a high proportion considering all other
activities such as control of weeds, harvest, and transportation. It should be considered,
Table 3
Dry matter production of ‘Clone 20’ (Opuntia ficus-indica Mill.) at different sites and P fertilization in north-east
Brazil
Site P fertilization (kg P ha1 yr1) P levela
0 33
1
t DM ha harvest1
Arcoverde 17.9 abb 20.2 b o0.15

S.B. do Una 12.9 bc 14.1 c o0.46
Serra Talhada 11.2 c 14.2 c o0.06
Sertânia 21.4 a 29.2 a o0.0001
S.E. ¼ 2.3 t DM ha1 harvest1.

a
b
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however, that in the first harvest the primary cladodes were not harvested, reducing in this
way the calculated N-use efficiency. In terms of P fertilization in the plant population of
40,000 plants ha1, each kg of added P yielded an average of 211 kg of DM. Considering
the same cactus price mentioned before (US$ 0.09 kg1) and the P price as US$ 1.00 kg1,
the P fertilizer represented only 5% of the selling price for the additional forage harvested.
Thus, adoption of P fertilization at this level (33 kg P ha1 yr1) involves less risk to the
farmers. It needs to be emphasized, however, that P response occurred only in Sertânia
(po0:0001) and marginally in Serra Talhada (po0:06), but not in the other two sites.
Therefore, soil P levels must be considered when doing these calculations. In the sites that
the responses were greater, soil P level was o10 mg kg1.
3.2. Total number of cladodes per plant
There was a triple interaction of site by plant population by P fertilization (Table 4).
Phosphorus fertilization increased total cladode number in Arcoverde and Sertânia for
both plant populations, but unlike DMY, cladode number did not increase with added P in
Serra Talhada. Although there is a positive correlation between total cladode number and
DM yield, other factors like total cladode weight and total cladode area are also positively
correlated with DM yield (Lucena et al., 1999). Sites did not differ in terms of cladode
number per plant either when no P was added in the 5000 plants ha1 plant population or
when P was added in the 40 000 plants ha1 (Table 4). Total cladode number per plant was
generally greater at the smaller plant population (5000 plants ha1) compared to
40 000 plants ha1 (Table 4). Because each individual plant in the 5000 plants ha1 density
had more area to explore, they had a greater development compared to the plants in the
higher population. On the other hand, the higher population had 8 times more plant per
unit area. Therefore, the total cladode number per area was higher in the higher plant
population (e.g., 6.75 and 26 cladodes per m2 in Sertânia for the 5000 and
40 000 plants ha1 with 33 kg ha1 of P, respectively). Because there is a positive correlation
between total cladode number and DM yield (Lucena et al., 1999), the productivity in the
higher plant population was higher. The primary cladodes were preserved at the harvest,
therefore, that unaccounted biomass leaded to the underestimation of the plant
productivity with the current data.
Table 4
Total number of cladodes per plant as affected by plant population, site and P fertilization
Site 5000 plants ha1 40 000 plants ha1
0 kg P ha1 yr1 33 kg P ha1 yr1 Pa 0 kg P ha1 yr1 33 kg P ha1 yr1 P
Cladode number plant1 Cladode number plant1
Arcoverde 6.4 ab 9.7 b o0.01 4.4 b 5.6 a 0.03

S.Talhada 7.0 a 7.7 c 0.25 6.5 a 6.4 a 0.83
Sertânia 7.9 a 13.5 a o0.01 4.8 ab 6.5 a 0.004
S.E. ¼ 0.7 cladodes plant1.

a
b
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Table 5
Total number of cladodes per plant as affected by plant population, and N and P fertilization
N level 5000 plants ha1 40 000 plants ha1
0 33 0 33
0 7.0 10.6 4.1 5.4

75 6.7 12.3 5.5 5.9
150 7.2 9.8 5.2 6.2
225 7.9 9.2 6.0 6.4
300 6.8 9.5 5.3 6.9
Effect (P level) NS (0.10) Linear (0.01) Quadratic (0.006) Linear (0.0002)
Standard error 2.1 2.0 1.0 0.8
Plant population also interacted with N and P fertilization affecting total cladode
number per plant (Table 5). Nitrogen fertilization had contrasting effect on total cladode
number. In the 5000 plants ha1 plant population, no N effect was observed without P
fertilization but when P was added (33 kg ha1 yr1 of P) a linear decrease in the total
cladode number as N fertilization increased from 0 to 300 kg ha1 yr1 of N was observed
(Table 5). In contrast, with 40 000 plants ha1 the total cladode number increased with N
fertilization showing a quadratic response at 0 P fertilization level and a positive linear
response at the level of 33 kg ha1 yr1 of P (Table 5). Increasing plant population
increased nutrient demand, which in turn modify the crop response to the N fertilizer.
Negative responses observed in the lower plant population was likely caused by a toxic
effect of the N fertilizer considering that the same amount of N was allocated for a
population 8-fold lower compared to the higher density (40 000 plants ha1).
3.3. Cladode area index
There was an interaction of plant population by site affecting the CAI as shown in Table
6. The CAI was greater at the higher plant population (40 000 plants ha1) for all evaluated
sites. Interaction occurred because at 5000 plants ha1, CAI in Sertânia and S.B. do Una
was similar, but at higher plant population (40 000 plants ha1) CAI was greater in
Sertânia (Table 6). Because the light interception is a critical step to increase plant growth,
increasing CAI leaded to greater plant productivity. In addition, fewer occurrences of
weeds are expected due to lower light levels reaching the soil in more dense plant
populations. Despite the increase in the CAI observed in the higher plant population, this
index reached up to 1, which still may be considered low in terms of light interception,
considering the upward position of the cladodes. Making a parallel with other forage
plants, Nascimento (1997) reported that legumes reach their growth potential at a lower
leaf area index (LAI) when compared to grasses due to differences in leaf architecture
(upward position in grasses and more horizontal position in legumes). According to Blaser
(1988), the optimum LAI for some grasses is in the 4–5 range. Therefore, a CAI close to
this value should also optimize spineless cactus productivity, considering the upward
position of the cladodes. In fact, Cortázar and Nobel (1986) reported that O. ficus-indica
productivity may increase when CAI (or SAI, stem area index) increase up to 4. Han and
Felker (1997) found increasing productivity of Opuntia ellisiana as the CAI increased from
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Table 6
Cladode area index of Clone 20 as affected by plant population and site in north-east Brazil
Cladode area index
S.B. do Una 0.28 ab

0.71 b o0.0001
Serra Talhada 0.02 b 0.20 c o0.0001
Sertânia 0.26 a 1.02 a o0.0001
S.E. ¼ 0.06.
a
b
0.06 (end of year 1) up to 3.88 (end of year 4). Despite this greater CAI reached in the 4th
yr (3.88) compared to CAI reached in the Brazilian research (1.0), the productivity
observed for O. ellisiana was lower and that may be explained by the species difference,
with O. ficus-indica being considered a faster growing cactus than O. ellisiana (Han and
Felker, 1997). Mizrahi et al. (1997) reported productivity of 40–50 t ha1 yr1 of DM for
spineless cactus with 240 000 plants ha1 without water or soil fertility restrictions. This
level of productivity (40 t DM ha1 yr1) was also confirmed by Cortázar and Nobel (1991)
who reported that the DM productivity of O. ficus-indica could reach 50 Mg ha1 yr1
based on their prediction model. It must be considered, however, that in north-east Brazil
the spineless cactus is grown only in rain fed systems. Despite of that, more studies aiming
to reach the potential of this crop in the prevailing conditions of the semi-arid region is still
necessary. The smallest CAI occurred in Serra Talhada for both plant populations (Table
6). The lower altitude in Serra Talhada (429 m above sea level) when compared to the other
sites (558 m in Sertânia, 614 m in S.B. do Una, and 663 m in Arcoverde) may be a possible
explanation for less productivity of the clone 20 in that location, considering that rainfall
was similar to the other sites. In the tropics, higher altitudes are associated to lower night
temperature, increasing the air relative humidity at night, which is ideal for the Opuntia
development due to its CAM physiology, particularly in semi-arid regions where water
deficit occurs to a greater extent (Sampaio, 2005).
CAI was also affected by an interaction of site by P fertilization with CAI increasing
with P fertilization only in Sertânia (Table 7). Because among the sites with lower P
levels (Serra Talhada and Sertânia, with 7 and 9 mg kg1 of soil P, respectively)
Sertânia presented higher productivity potential for Opuntia development, the response
was greater in that site when compared to Serra Talhada. Arcoverde and S.B. do Una
presented higher soil P levels and that is likely the reason for no response to added P in
terms of CAI.
There was also an interaction of plant population by P fertilization affecting CAI
(Table 8). The CAI was greater in the 40 000 plants ha1 population either with or without
P. Interaction occurred because the CAI increased with added P only at the highest plant
population with no increase with 5000 plants ha1 (Table 8). The same explanation for this
interaction in the other response variables holds for CAI, i.e. P demand is greater in higher
plant population increasing crop response to P fertilizer.
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Table 7
Cladode area index of Clone 20 as affected by site and P fertilization
Site P fertilization (kg ha1 yr1 of P) P levela
0 33
Cladode area index

b
S.B. do Una 0.49 a 0.50 b 0.68
Serra Talhada 0.11 b 0.11 c 0.97
Sertânia 0.51 a 0.77 a o0.0001
S.E. ¼ 0.06.
a
b
Table 8
Cladode area index of Clone 20 as affected by plant population and P fertilization
Plant population P fertilization (kg ha1 yr1 of P) P levela

(plants ha1)
0 33
Cladode area index
5000 0.17 bb 0.21 b 0.14

40 000 0.57 a 0.71 a o0.0001
S.E. ¼ 0.04.
a
b
3.4. Rainfall use efficiency (RUE)
Because rainfall was different among the four evaluated sites, it is important to analyse
the RUE in order to standardize the results. In general, improving crop management
increased RUE, i.e. more DM produced per unit of rainfall. The average RUE for all
treatments and sites was 18 kg DM ha1 mm1 of rainfall, i.e. 555 kg H2O kg1 DM
(SE ¼ 11.8 kg DM ha1 mm1 of rainfall). This value did not include the primary cladodes
because they were not harvested, thus, the RUE was underestimated for all sites. In
addition to that, the water losses through evaporation, surface runoff, and deep drainage
were not taken in account, contributing to the underestimation of the water use efficiency.
Han and Felker (1997) in a long-term field evaluation found a water-use efficiency of
162 kg H2O kg1 DM for O. ellisiana. These authors, however, estimated the water balance
including evapo-transpiration, runoff and, deep drainage. Significant interactions occurred
between site and plant population (Table 9). In all the sites, the higher RUE occurred for
the higher plant population. Interaction occurred between site and plant population
because at the lower plant population only Serra Talhada differed from the other sites
presenting lower RUE. At the higher plant population, however, Arcoverde and Sertânia
showed greater RUE, followed by São Bento do Una and Serra Talhada. Increased plant
ARTICLE IN PRESS
Table 9
Rainfall use efficiency (kg DM ha1 mm1 of rainfall) of ‘Clone 20’ (Opuntia ficus-indica Mill.) at different sites
and plant populations in north-east Brazil
kg DM ha1 mm1of rainfallb
Arcoverde 13.1 ac 23.0 a o0.0001

S.B. do Una 11.5 a 22.1 bc o0.0001
Serra Talhada 5.0 b 16.1 c o0.0001
Sertânia 17.6 a 34.9 a o0.0001
S.E. ¼ 2.3 kg DM ha1 mm1 of rainfall.

a
b
Primary cladodes not included in the calculation because they were not harvested.
c
population resulted in greater RUE for all four locations. More plants per unit area
increased the light interception and the cactus growth per area, reducing the weeds leading
to greater efficiency of water utilization. Turner (2004) reported that agronomic practices
that reduce water losses and improves crop yield increases RUE. Agronomic procedures
such as minimum tillage, appropriate fertilizer use, improved weed/disease/insect control,
timely planting, and a range of rotation options are example of practices to increase RUE.
Thus, increasing plant population, N, and P fertilization increased RUE of spineless cactus
because of greater DM harvested with the same rainfall. Serra Talhada showed the greatest
rainfall (1204.4 mm) during the growth period and São Bento do Una the least (805.8 mm).
Because the DM harvested was similar for both locations (Table 3), the RUE was greater
in São Bento do Una due to less rainfall. Rainfall distribution may also affect RUE
because of water losses through runoff and leaching. In general, rainfall distribution was
more uniform in São Bento do Una than in Serra Talhada. Different soil orders (Entisol in
São Bento do Una and Alfisol in Serra Talhada) probably affected RUE as well. Different
sand and clay proportions in the soil affects the soil water holding capacity and soil
available water. Usually, a clayey soil has greater soil water holding capacity and soil
available water, however, the clayey soil holds the water to a greater tension at the same
soil water-content compared to a sandy soil (Hillel, 1998). Thus, the plant is able to
absorb water at lower soil water-content in a sandy soil compared to a clayey soil, affecting
the RUE.
There was also an interaction between site and P fertilization (Table 10). This interaction
occurred because Sertânia was the only site that showed greater RUE with increased P
fertilization. Because the greater response to P in Sertânia, RUE was also greater in
Sertânia for the higher P fertilization. The low soil P level observed in Sertânia (9 mg kg1)
may explain this result. Despite the low soil P level also observed in São Bento do Una
(7 mg kg1), the RUE was not affected by P fertilization (p ¼ 0:12). Plant population also
interacted with N fertilization. In this case, there was no significant effect of N fertilization
at the lower plant population, but there was a linear increase for RUE with increased N
levels when N was applied at the higher plant population. Finally, plant population
ARTICLE IN PRESS
Table 10
Rainfall use efficiency (kg DM ha1 mm1 of rainfall) of ‘Clone 20’ (Opuntia ficus-indica Mill.) at different sites
and P fertilization in north-east Brazil
Site P fertilization (kg P ha1 yr1) P levela
0 33
kg DM ha1 mm1of rainfall
Arcoverde 16.9 ab 19.1 b 0.17

S.B. do Una 16.1 a 17.5 bc 0.36
Serra Talhada 9.3 b 11.8 c 0.12
Sertânia 22.2 a 30.3 a o0.0001

a
b
Table 11
Rainfall use efficiency (kg DM ha1 mm1 of rainfall) of ‘Clone 20’ (Opuntia ficus-indica Mill.) at different plant
populations and P fertilization in north-east Brazil
Plant population P fertilization (kg P ha1 yr1) P levela
0 33
kg DM ha1 mm1of rainfall
5000 pl. ha1 10.9 bb 12.7 b 0.11

40,000 pl. ha1 21.4 a 26.6 a o0.0001

a
b
interacted with P fertilization (Table 11), with no P response at 5000 plants ha1. Response
to P fertilization occurred only at the higher population. Greater P demand at higher plant
population may be part of the explanation. Similar response was observed for the N
fertilization, i.e. N application only affected RUE at the higher plant population.
3.5. Plant nitrogen concentration
There was a site by N fertilization interaction affecting N concentration in the forage

(Fig. 1). A positive linear effect of N fertilization on plant N concentration was observed in
S.B. do Una, Arcoverde, and Serra Talhada, and a quadratic effect was observed in
Sertânia (Fig. 1). The N concentration was within the range of 6.7 g kg1 (Serra Talhada;
0 kg N ha1 yr1) to 13.9 g kg1 (Arcoverde; 300 kg N ha1 yr1). The increase in plant N
concentration with N fertilization of Opuntia is reported in the literature. Gonzalez (1989)
observed that crude protein levels of O. lindheimeri Engelm. were increased by 3.1, 4.2,
and 4.4 percentage units by the application of 67, 135, and 224 kg N ha1, respectively.
ARTICLE IN PRESS
Arcoverde
16
y = 0.02x + 7
R2 = 0.95;P < 0.001
14
S. B. do Una
12 y = 0.007x+10.8
R2 = 0.67; P < 0.03
10 Serra Talhada
y = 0.02x+ 6.8
N, gkg-1
8 R2 = 0.99;P < 0.001
Sertânia
6 y = 4E-05x2 - 0.009x + 8.3
Arcoverde R2 = 0.70; P < 0.05
4 S.B. do Una
Serra Talhada
Sertânia
2
0
0 50 100 150 200 250 300
kg N ha-1 yr-1
Fig. 1. Nitrogen concentration in Opuntia ficus-indica cv. Clone 20 as affected by N fertilization and location in
north-east Brazil.
14
12 5000 plants/ha
40000 plants/ha
Plant N concentration, g kg-1
10
8
a
6
b
0
5000 plants/ha 40000 plants/ha
Plant population
Fig. 2. Nitrogen concentration in Opuntia ficus-indica cv. Clone 20 as affected by plant population in north-east
Brazil.
ARTICLE IN PRESS
In the current research, linear responses occurred in three out of four locations.
Quadratic response occurred only in Sertânia where plant N concentration decreased
with initial N level (75 kg N ha1) increasing thereafter. Because Sertânia was the site
with the greatest Opuntia development, the dilution effect may be part of the explanation
for the quadratic response at initial N levels. Minson (1990) suggested that forage
crude protein (CP) concentration should be at least 70 g kg1 (i.e. 11.2 g N kg1) for
normal rumen functioning. In the current research, this level was reached only with N
fertilization. Other way to overcome the CP deficiency for the ruminant fed with
Opuntia is the supplementation with protein concentrate or even non-proteic N (Melo
et al., 2003).
Plant population also affected N in the forage (Fig. 2), with greater plant N
concentration observed in the population of 5000 plants ha1 (11.2 g kg1; SE ¼ 0.3 g kg1)
compared to the population of 40 000 plants ha1 (9.0 g kg1). Individual plant develop-
ment was greater in the lower plant population and the number of cladodes was also higher
(Table 4). Because the new cladodes have greater N concentration than the older ones
(Santos et al., 1990a), the greater number of them results in a higher average N
concentration in the whole plant, which occurred in the 5000 plants ha1 population. Nerd
and Nobel (1995) reported higher nitrate levels and higher nitrate reductase activity in new
cladodes compared to the older underlying ones. They also found higher nitrate levels in
the inner water-storing layer (parenchyma) than in the outer green photosynthetic layer
(chlorenchyma).
4. Conclusions
Nitrogen fertilization increased productivity of O. ficus-indica cv. ‘Clone 20’ only at the
higher plant population, i.e. 40 000 plants ha1. Plant N concentration also increased with
N fertilization but the degree of response varied with site.
Phosphorus fertilization increased productivity of O. ficus-indica cv. ‘Clone 20’ only
when soil P levels were o10 mg kg1.
Greater plant population is necessary to achieve higher productivity. Because of the low
CAI obtained at the harvest time, there is a potential to increase the DM yield of O. ficus-
indica cv. ‘Clone 20’ by increasing the population beyond the limit used in this research, i.e.
40 000 plants ha1. Chemical fertilization and more dense plant population improved RUE
resulting in more forage produced per unit of rainfall.
Acknowledgments
Thanks to World Bank which provided partial financial assistance for this project
through the PRODETAB/EMBRAPA program.
Appendix A. Supplementary materials
Supplementary data associated with this article can be found in the online version at
doi:10.1016/j.jaridenv.2006.02.015.
Appendix B. Supplementary materials

ARTICLE IN PRESS
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Productivity of Opuntia ficus-indica (L.) Miller

Corresponding author. Tel.: +55 81 33206574.

Keywords: Cactaceae; CAM; Semi-arid; Spacing; Spineless cactus

2. Material and methods

Site Planting date Harvest date

São Bento do Una May 12, 1998 April 2000

3. Results and discussion

3.1. Dry matter yield

Site Plant population P levela

5000 plants ha1 40 000 plants ha1

Arcoverde 13.8 abb 24.3 b o0.0001

S.E. ¼ 2.3 t DM ha1 harvest1.

Site P fertilization (kg P ha1 yr1) P levela

Arcoverde 17.9 abb 20.2 b o0.15

S.E. ¼ 2.3 t DM ha1 harvest1.

3.2. Total number of cladodes per plant

Site 5000 plants ha1 40 000 plants ha1

0 kg P ha1 yr1 33 kg P ha1 yr1 Pa 0 kg P ha1 yr1 33 kg P ha1 yr1 P

Cladode number plant1 Cladode number plant1

Arcoverde 6.4 ab 9.7 b o0.01 4.4 b 5.6 a 0.03

S.E. ¼ 0.7 cladodes plant1.

N level 5000 plants ha1 40 000 plants ha1

0 7.0 10.6 4.1 5.4

3.3. Cladode area index

Site Plant population P levela

5000 plants ha1 40 000 plants ha1

Cladode area index

S.B. do Una 0.28 ab

Site P fertilization (kg ha1 yr1 of P) P levela

Cladode area index

Plant population P fertilization (kg ha1 yr1 of P) P levela

Cladode area index

5000 0.17 bb 0.21 b 0.14

3.4. Rainfall use efficiency (RUE)

Site Plant population P levela

5000 plants ha1 40 000 plants ha1

kg DM ha1 mm1of rainfallb

Arcoverde 13.1 ac 23.0 a o0.0001

S.E. ¼ 2.3 kg DM ha1 mm1 of rainfall.

Site P fertilization (kg P ha1 yr1) P levela

kg DM ha1 mm1of rainfall

Arcoverde 16.9 ab 19.1 b 0.17

S.E. ¼ 2.3 kg DM ha1 mm1 of rainfall.

Plant population P fertilization (kg P ha1 yr1) P levela

kg DM ha1 mm1of rainfall

5000 pl. ha1 10.9 bb 12.7 b 0.11

S.E. ¼ 1.3 kg DM ha1 mm1 of rainfall.

3.5. Plant nitrogen concentration

There was a site by N fertilization interaction affecting N concentration in the forage

8 R2 = 0.99;P < 0.001

Appendix A. Supplementary materials

Appendix B. Supplementary materials

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