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Flowering levels, harvest season and yields of cupuassu (Theobroma grandiflorum)

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Cupuassu (Theobroma grandiflorum), a specie native to Amazonia, has been planted commercially in Brazil to satisfy the demand for the flavorful juice obtained from the pulp around its seeds. The trees are notorious for low and irregular fruit production. Data gathered over two seasons from trees in a germplasm collection in Pará, Brazil, showed that some of them fruited more regularly than others. Differences in fruit production correlated to differences in flower production. Tree-to-tree variation in flower production, fruit production, and consistency of both over time suggest considerable scope for improving yields by selection. Hand pollinations resulted in a much higher frequency of fruit set than open pollinations, indicating that lack of effective pollination is also a reason for low yield. However, attempts to increase the level of effective pollination are handicapped by low knowledge about the pollinators of cupuassu and their behavior.
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143 VOL. 41(1) 2011: 143 - 152
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiorum)1
Giorgini Augusto VENTURIERI1
ABSTRACT
Cupuassu (eobroma grandiorum), a specie native to Amazonia, has been planted commercially in Brazil to satisfy the
demand for the avorful juice obtained from the pulp around its seeds. e trees are notorious for low and irregular fruit
production. Data gathered over two seasons from trees in a germplasm collection in Pará, Brazil, showed that some of them
fruited more regularly than others. Dierences in fruit production correlated to dierences in ower production. Tree-to-tree
variation in ower production, fruit production, and consistency of both over time suggest considerable scope for improving
yields by selection. Hand pollinations resulted in a much higher frequency of fruit set than open pollinations, indicating that
lack of eective pollination is also a reason for low yield. However, attempts to increase the level of eective pollination are
handicapped by low knowledge about the pollinators of cupuassu and their behavior.
KEYWORDS: eective pollinator, ower production, fruit set, irregular bearing
Níveis de oração, período de safra e produção do cupuaçuzeiro
(Theobroma grandiorum)
RESUMO
Cupuaçu (eobroma grandiorum) é uma espécie nativa da Amazônia que vem sendo plantada comercialmente no Brasil para
satisfazer a demanda pelo seu suco, de sabor marcante, obtido da polpa que envolve as suas sementes. As árvores não notórias
pela sua baixa e irregular produtividade. Dados obtidos de dois períodos produtivos, de árvores de uma coleção de germoplasma
no estado do Pará, Brasil, mostraram que algumas delas fruticam mais irregularmente que outras. Diferenças na produção de
frutos foram correlacionadas à diferenças na produção de ores. Variações entre plantas nas produções de ores e de frutos, e a
repetibilidade de ambos ao longo do tempo sugerem melhoria das colheitas por seleção. Polinizações manuais resultaram em
aumento no pegamento dos frutos, indicando que a falta de polinizador efetivo é também um causa da produtividade baixa.
Entretanto, tentativas para aumentar o nível de polinização efetiva são dicultadas pelo pouco conhecimento sobre os agentes
polinizadores do cupuaçu e seu comportamento.
PALAVRAS-CHAVE: polinizador efetivo, produção de ores, pegamento dos frutos, colheitas irregulares.
1Universidade Federal de Santa Catarina; Email: giorgini@cca.ufsc.br
1 The present paper is part of the PhD thesis of the author.
144 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
INTRODUCTION
Cupuassu (eobroma grandiorum (Willd. ex Sprengel)
Schumann, family Sterculiaceae) is a native of eastern
Amazonia. Wild trees occur in the underst orey of the forests
of eastern Pará and western Maranhão in Brazil. Cupuassu is
planted and protected by various indigenous groups and is
cultivated in house gardens as far west as Peru and Colombia
by non-indigenous settlers (Smith et al. 1992). As in its
better-known relative, the cacao (T. cacao L.), the seeds are
rich in fat and are surrounded by a juicy pulp, but the fat of
cupuassu seeds has a lower melting point than that of cacao
(Silva 1988), so cupuassu is not generally used in chocolate
manufacture. However, the characteristic acid-sweet avor
of the pulp has proved widely acceptable both within Brazil
and internationally and cupuassu is expanding rapidly into
the world market for exotic fruit avors for use in juices and
ice-creams (Clement and Venturieri 1990). By the 1980s
demand exceeded supply and cupuassu was one of the most
expensive local fruits in Amazonia. A single fruit would sell for
up to US$2.00, and a tonne of frozen pulp fetched US$4,000
(Venturieri 1993). is has resulted in a great increase in
cupuassu plantations. Cupuassu is also often recommended as
a cash crop for multi-crop integrated agroforestry systems, in
part because it is potentially so protable and in part because,
as a member of the forest understorey, it is tolerant of shade
(Moraes et al. 1994; Smith et al. 1992; Venturieri 1993).
e fruits of cupuassu are the largest known in eobroma.
ey weigh 200-4,000 g and contain 30-50 seeds, which
account for 17% of the fruit weight. Relatively few owers
develop into fruits. Yields are low compared to those of other
fruit trees and yields per tree are extremely variable (Venturieri
1993). e reasons for this have not yet been established, but
may relate to inadequate pollination, inadequate supply of
nutrients, or both. Like other species of eobroma, cupuassu
has structurally complex owers. A biotic vector of some kind
is required to convey pollen from the petal pouches, into which
it is shed, to the stigmas. Most trees are self-incompatible
(Venturieri 1994), so the pollinator must move between trees
to bring about eective pollination, though there are some
reports of isolated trees which produce fruit, presumably by
self-pollination (Venturieri 1993). Venturieri (1994) found
that fewer than 2% of naturally-pollinated cupuassu owers
had over 50 pollen grains on their stigmas. is is low even
for eobroma, in which only 2-4% of owers are recorded
as being eectively pollinated in T. cacao (Free 1993; Parvais
et al. 1977), 7.5% in T. obovatum Klotzsch ex Bernouilli
(Souza and Venturieri 1998), 18.7% in T. subincanum Mart.
(Rodrigues and Venturieri 1997) and 20% in T. speciosum
Willd. ex Sprengel (Souza and Venturieri, 2010). Selecting
for more owers per tree might increase yield by increasing
the attractiveness of the tree to the pollinator (Faegri and van
der Pijl 1979). Changes in the length of the owering season,
particularly insofar as these increase or decrease competition
with other plant species which use the same pollinators, or
extend the period during which particular pollinators can
remain active within the plant community as a whole, might
also aect the proportion of owers which are eectively
pollinated and hence aect yield. Pollen vectors are very
dicult to identify in eobroma, but in cupuassu the most
likely candidates are chrysomelid weevils (Venturieri et al.
1997) and stingless bees (Venturieri 1994). Small pollinators
such as these may not y freely during periods of heavy rain, so
weather during the owering season may also inuence yield.
Species with fruits which are energetically expensive to
produce, such as cacao or avocado (Persea americana L.),
often mature relatively few fruits compared to the numbers
of owers produced because the mother tree is physiologically
incapable of lling all the fruits which set. In this case, fruits
are aborted after pollination, not because of a failure of
pollination (Stephenson 1981), and improving the nutrient
status of the trees may increase the number of fruits which
mature successfully.
Commercial production of cupuassu requires the fruits
to be processed to extract the juice from the pulp before the
pulp ferments, and the extracted juice then has to be frozen
or pasteurized. Processing plants depend on a predictable and
even workload throughout the year to keep the machinery and
labor force operating at optimum capacity. It is important to
be able to forecast cupuassu yields as early as possible each
season for this reason and also so that growers may warn of
possible problems in fullling contracts.
Little information is available on variation within cupuassu
with respect to owering period, rate at which fruits mature,
relation between number of owers produced and yield of
fruit, eects of weather conditions at or after pollination,
regularity of bearing, and response to irrigation or articial
fertilizers. is information is needed in order to assess
whether selection for these features is likely to be an eective
strategy for improvement of cupuassu, either as a plantation
crop or as a cash crop for cooperatives of small-scale farmers
cultivating without totally clearing the rain forest. is study
provides preliminary data on some of these points.
MATERIALS AND METHODS
Study population
e trees used in this study included 20 from the Basil
Bartley living collection of species of eobroma and 94 from
a sub-spontaneous population of mature cupuassu trees. e
trees were maintained at the “José Haroldo” Experimental
Station for Cocoa Genetic Resources, of the Executive
Commission for Cocoa Crop Planning (CEPLAC), in the
145 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
municipality of Marituba, Pará, Brazil (1°12’S; 49°13’W).
e trees in the Basil Bartley collection were planted in 1984
from seeds of three dierent fruits, collected in dierent
parts of Amazonia. eir identifying numbers are preceded
by the letter F. e origin of the sub-spontaneous trees is not
recorded, but they are thought to derive from trees planted
by the caboclos (peasant farmers) who originally owned the
land on which the Experimental Station now stands. eir
ages have been roughly estimated from the sizes of their trunks
as ranging from ten to 85 years (Venturieri 1994).
All trees were cultivated on the type of oxisol common
in the uplands of the state of Pará, with limited drainage and
low fertility. For three years prior to the start of this study,
trees in the Basil Bartley collection had been treated annually
with 0.15 kg of articial fertilizer (NPK 10-30-20 formulated
using ingredients produced by Ultrafertil S/A – Catalão - GO),
while the sub-spontaneous trees were not treated. In December
1991, seven of the study trees (three subspontaneous trees and
four trees in the Basil Bartley collection) received a mixture of
equal parts of articial fertilizer and Yoorin MG (produced by
Fertilizantes Mitsui S/A- Poços de Caldas - MG, containing
P2O5 18%, Mg 14.5%; CaO 28%, and traces of Zn, Bo, Si,
Fe, Mn, Co and Cu). e trees in the Basil Bartley collection
received 0.3 kg of this mixture, the older sub-spontaneous
trees received 1 kg. e subspontaneous trees received more
fertilizer because the trees were bigger.
e monthly rainfall data during the period of the study
were available from the records made by the Experimental
Station.
Length of owering season and number of owers
produced
Observations were made from June to December 1991 and
from May 1992 to January 1993.e number of owering
trees was recorded each 15 days and the trees were classied
on the basis of the number of open owers per tree: zero = no
open owers; one = up to ten open owers; two = 11-50 open
owers and three = more than 50 open owers. e length of
the owering season for each tree was considered as the time
it bore any owers.
e number of owers produced per tree was calculated
from the number of 15-day periods the tree spent in each of
the three classes before mentioned, multiplied by the mean
number of owers for that class (considered as 70 owers for
class three). Since each ower lasts only one day, each class
gure so obtained was multiplied by 15, and the totals for the
three classes then summed as an estimate of the total number
of owers produced by each tree.
Fruit production, time to maturity and length of
harvest season
e numbers of fruits produced by open pollination in
1991/92 and 1992/93 (hereafter referred to as rst and second
season, respectively) were counted on each tree on December
30th 1991 and 1992, after they had passed the critical period
for fruit drop but before they were fully mature
During the second season, some owers on eight trees
were hand pollinated, as described by Venturieri and Ribeiro
Filho (1995). e time to fruit maturity, i.e. number of days
from pollination to fruit fall, was calculated for these fruits
because their dates of pollination were known with certainty.
e fruits were enclosed in net-bags and checked for fallen
fruit each 15days during both seasons in order to assess the
length of the fruiting season.
Statiscal analyses
Correlations among days from pollination and weight
of fruit, length and intensity of owering, total number of
owers and total number of fruits per tree in each season were
assessed by Pearson’s correlation coecient (Sokal and Rohlf,
1995). Number of days from pollination to fruit ripening were
statistically compared using t test (Sokal and Rohlf, 1995).
RESULTS
Phenology of owering
Comparison of monthly rainfall (Figure 1) with the
percentage of trees owering each month (Figure 2) shows
that, in both rst and the second season, peak owering
occurred during the driest period of the year (July-September
in 1991; October-December in 1992). e fact that maximum
owering in this population can vary by as much as three
months suggests that owering is not controlled by daylength,
which in any case varies very little at the latitude of this study.
Water supply for the trees is never limiting since there is
never a soil water decit (Garcia et al. 1985). Light intensity
is markedly lower during the rainy season because of cloud
cover, and light intensity may be the major factor controlling
initiation of owers in cupuassu, as has been suggested for
Hevea brasiliensis, another amazonian species, that owering
months generally coincide with the months of the highest
average solar irradiation measured at the ground not in the
upper atmosphere (Hoong-Yeet 2007).
In the rst season, 68% of the trees owered simultaneously
in the second half of August, and in the second season, 55%
of the trees owered simultaneously in the second half of
November (Figure 2). Over half of the trees in this population
were therefore potentially able to inter-pollinate.
146 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
In the rst season, ten (8.8%) of the trees studied never
owered, while in the second season 31(27.6%) failed to
ower. Eight (7.14%) trees did not ower in either season.
Some trees owered very sparsely, and for a limited period only
(e.g. tree 200-26, Figure 3a). Other trees owered abundantly,
either in both seasons (e.g. trees 229-25 and F13-2, Figure
3b) or in one season only (e.g. tree 513-23 in the rst season,
Figure 3b). Trees that owered abundantly produced most
of their owers in a short period of peak owering (class
3), usually preceded and followed by periods of less intense
owering (class 1 or class 2).
Phenology of fruiting
In the rst season 178 ripe fruits were produced from
open pollination, but only 48% (50 out of 104) of the trees
which owered in that season. Mean number of fruits per
tree, considering only the fruiting trees, was 6.6. In the second
season, 162 fruits were produced by 44% (35 out of 79) of
the trees which owered, with a mean of 4.5 fruits per fruiting
tree. Only 55 out of over 100 trees studied produced any fruit
during the period of these observations.
e numbers of fruits set on the most productive and
least productive trees in each season are shown in Table 1
is clearly illustrates the great variation in yield between
dierent trees in the same season (from 0 to 28 fruits per
tree) and the same tree in dierent seasons (from eight to 28
fruits). e data suggest that a tree which fruits heavily in
one season (e.g. trees 513-23 and 492-28) may set relatively
few fruit in the following season, but unfortunately the
observations could not be continued for long enough to
establish whether there is really a tendency to biennial or
irregular bearing in this species.
e percentage of owers which set fruit following hand
pollination was much greater than the percentage which set
fruit following natural pollination (10% versus less than
0.5%). is suggests that eective pollination may be the
Most productive trees Least productive trees Mean
Number of fruits set Number of fruits set
Tree First season Second season Total Mean Tree First season Second season Total
229-25** 1 36 (22) 37 18.5 200-26 1 0 1 0.5
492-60** 6 31 (6) 37 18.5 216-23 0 1 1 0.5
513-23 28 8 36 18 223-34 1 0 1 0.5
492-28 26 4 30 15 288-22 0 1 1 0.5
283-56 7 23 (19) 30 15 298-19 1 0 1 0.5
545-33 8 21 29 14.5 302-36 1 0 1 0.5
534-33 19 10 29 14.5 321-43 0 1 1 0.5
296-1 2 24 (12) 26 13 498-3 1 0 1 0.5
332-17 5 15 (9) 20 10 F13-8* 1 0 1 0.5
F13-2* 7 13 (4) 20 10 Various trees 1 1 2 1
Mean 10.9 18.5 Mean 0.7 0.4
Table 1 - Fruit production in consecutive seasons on some individual trees of cupuassu (figures in parentheses = fruits set by hand pollination; * tree treated
with 0.3 kg fertilizer, ** tree treated with 1.0 kg fertilizer).
Figure 1 - Monthly rainfall at the Experimental Station for Cocoa Genetic Resources “José Haroldo”(ERJOH), Marituba, Pará, Brazil during the period of study
(* = data not available).
147 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
variation in the timing of the preceding owering seasons
(Figure 2).
Correlations between length and intensity of
owering, total ower production, and fruit
production
Table 3 shows the correlations between all pairs of these
variables. Not surprisingly, the total number of flowers
produced was correlated more strongly with the lengths of the
periods of intense owering (classes 2 and 3) than with the
total length of the owering season, which included the period
of sparse (class 1) owering. In both seasons, the number of
Female tree N Days from pollination to fruit ripening
(in average by female tree)
338-33 2 137.5a
339-42 3 151.3ab
332-17 9 159.9ab
F13-2 4 161.2ab
229-25 22 165.3 b
492-60 6 166.3 b
283-56 19 167.5 b
296-1 12 171.6 b
Mean 164.8
Minimum 105
Maximum 188
Table 2 - Number of days from pollination to fruit ripening of 77 fruits from 8 female cupuassu (Theobroma grandiflorum) trees. Averages followed by the same
letter are not significantly different according to the t-test.
principal factor limiting fruit set, not nutrient status or
physiological capacity of the trees.
Mean time from pollination to fruit maturity for the
77 fruits obtained from hand-pollination was 165 days
(minimum 105, maximum 188), or approximately 5.5
months (Table 2). e correlation analysis between weight
of fruit and days from pollination to their ripening was very
low and not signicant (r= 0.19 p= 0.1). So, some trees
mature their fruits faster than others. In both seasons, most
fruits set by open pollination ripened between December
and May, but the precise months in which ripe fruits were
most abundant varied from year to year (Figure 4), reecting
Length of period in flowering class Total length of
flowering period
Total no. of flowers
per tree
1
(1-10 flowers)
2
(11-50 flowers)
3
(>50 flowers)
Length of period in flowering class
2 (11-50 flowers)
-0.14
0.29**
Length of period in flowering class
3 (>50 flowers)
0.00
0.21
0.13
0.76**
Total length of flowering period 0.64**
0.87**
0.63**
0.72**
0.36**
0.61**
Total no. of flowers per tree 0.02
0.41**
0.76**
0.93**
0.73**
0.93**
0.75**
0.80**
Total no. of fruits per tree -0.02
0.43**
0.12
0.60**
0.50**
0.73**
0.20
0.66**
0.40**
0.74**
Table 3- Correlation coefficients between length of flowering, total number of flowers per tree and fruit production per tree (upper figure in each cell relates to
the first season, lower figure to the second season; ** significant at the 1% level).
148 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
fruits produced per tree was signicantly correlated with the
total number of owers per tree.
DISCUSSION
Fewer than 50% of the trees observed over two
consecutive seasons produced any fruit. e mean number
of fruits set per tree, from open pollination, was appreciably
less (4.5-6.6) than the 12 fruits per tree obtainable from
well-managed plantings of cupuassu (Calzavara et al. 1984;
Falcão and Lleras 1983), and was comparable to the number
of fruit set on wild trees (Calzavara et al. 1984). ). Trees which
produced many owers and fruits in one season produced
only half or fewer of those in the following season (Figure
3b). Other trees yielded more evenly over consecutive
seasons, but their yields were appreciably below those of the
most productive trees.
The number of flowers produced per tree varied
between dierent trees in the same season and between
the same tree in dierent seasons in much the same way as
fruit production. Fruit production per tree is likely to be
determined partly by genotype, but also by external factors
such as compatibility with near neighbors, amount of light
received, and whether the tree is healthy or diseased. e
heavily-cropping trees may tend to irregular bearing, as in
many other fruit trees (Sedgley and Grin 1989). Our data
suggest that selection for yield and/or regular bearing may
be eective in this relatively unimproved crop, but more
carefully designed experiments, continued over a longer
period, are necessary to demonstrate this conclusively.
Venturieri (1994) found that two-thirds of the owers
produced by cupuassu remain unpollinated. is, combined
to the signicant number of fruits produced by hand-
pollination, suggests that the lack of eective pollination
may explain, in part, the limited fruit set. Fruit production
over the two seasons of our study was correlated slightly more
strongly with the length of the period of peak owering than
with the total length of the owering period or the total
number of owers per tree. is might suggest that trees
bearing more owers at a given time attract more pollinators,
as Warren et al. (1995) found in eobroma cacao. However,
production of more owers per day per individual is likely to
result in more geitonogamous pollinations which, in a self-
incompatible species like cupuassu (Venturieri 1994), would
be ineective.
Cupuassu owers probably oer three dierent rewards to
their insect visitors. Pollen is available as soon as the owers
Figure 4 - Period of fruit ripening in cupuassu trees studied (based on 178 fruits in the first season and 162 fruits in the second season, all set by open pollination).
Figure 2 - Flowering period and level of flowering in the total population of cupuassu trees studied.
149 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
Figure 3 - (a) Flowering period and level of flowering in some unproductive trees of cupuassu.(b) Flowering period and level of flowering in some productive trees of cupuassu.* = tree treated with 0.3 kg
fertiliser, ** = tree treated with 1.0 kg fertiliser.
ab
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Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
open, as is shelter in the petal pouches and/or within the
ring of staminodes. e copious stigmatic exudate is another
possible food reward. e owers usually open mid to late
afternoon and remain open for 24-36 hours, but the stigmatic
exudate is produced most abundantly between 2:00 and 6:00
hours (Venturieri 1994). Cantharophily (beetle pollination)
and Mellithophily (bee pollination) are accepted as possible
pollination syndromes in cupuassu (Venturieri et al. 1997,
Venturieri 1994). Regarding the two potential pollinators
so far identied, the chrysomelid weevils may shelter in
cupuassu flowers overnight, feed on pollen in the petal
pouches, and then move on the following day when the ower
wilts (Venturieri et al. 1997). It is not known whether these
weevils would initially be attracted preferentially to trees with
a larger display of owers, or whether they would then tend
to move from ower to ower on the same tree. Stingless
bees, the other potential pollinators (Venturieri 1994) are
more likely than weevils to be density-dependent foragers,
and thus likely to bring about ineective pollinations on self-
incompatible trees displaying many owers. However, these
bees are strictly diurnal, and they are most active when the
stigmas of cupuassu are not receptive (Venturieri 1994). eir
importance as eective pollinators of cupuassu has still to be
established. Venturieri (1994) found that the proportion of
cupuassu owers pollinated remained more or less constant
throughout the owering season. e correlation observed
here between length of the period of peak owering and fruit
production may therefore indicate only that a rare event, such
as fruit set in cupuassu, occurs more frequently as the number
of opportunities for it to occur increases.
Our evidence indicates that increasing the levels of eective
pollination in this population, and probably other populations
of cupuassu should signicantly increase the numbers of fruits
produced. However, until more is known about the relative
roles of the various potential pollinators, their life cycles and
their behavior, no recommendations can be made about how
to increase the proportion of owers pollinated.
ese preliminary and limited studies therefore suggest
that low and irregular yields are the characteristic feature of
cupuassu. It may nevertheless be possible to improve yield
signicantly in the short term by selection and in the longer
term by breeding, and also by further investigation of the
eects of factors such as provision or removal of shade, or
addition of fertilizer. Further studies on the pollination of
cupuassu may also be of practical as well as academic interest.
ACKNOWLEDGMENTS
is study was supported by a grant from the Margaret
Mee Amazon Trust and formed part of the Ph.D. thesis
conducted by G.A. Venturieri in the Department of
Agricultural Botany, e University of Reading, U.K. We
thank the Federal University of Pará (UFPa), the Executive
Commission for Cocoa Planning (CEPLAC), and Paraense
Museum Emílio Goeldi, for providing laboratory and eld
facilities necessary for this project. Our sincere thanks also
go to Sr. Edilson de Freitas Leal, of the CEPLAC technical
sta, for his assistance during the eldwork and Barbara
Pickersgill for suggestions given during the experiments and
contributions in the text.
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151 VOL. 41(1) 2011: 143 - 152 VENTURIERI
Flowering levels, harvest season and yields of
cupuassu (Theobroma grandiflorum)
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Recebido em 05/04/2010
Aceito em 07/07/2010
... The experiment was conducted over 12 years and followed a completely randomised design. This experimental design was necessary considering the negative effects of self-incompatibility and, in some cases, inter-incompatibility of the species (Venturieri, 2011). Thus, individuals of four different clones surrounded an individual of a given clone. ...
... Evaluations were conducted at the plant level over nine consecutive harvests (2010-2011 to 2018-2019). In the cupuassu tree, the harvest opening coincides with the beginning of the rainy season and extends over the entire period of about six months (Venturieri, 2011). Therefore, each harvest was divided into four evaluations with 45-day intervals between them. ...
... The grafting technique offers advantages, such as uniformity and early production, especially in perennial fruit trees such as T. grandiflorum (Baron, Amaro, Pina, & Ferreira, 2019). Fluctuations in production observed in this experiment during the stable productive phase are common in commercial crops and are attributed to climatic factors, especially long periods of drought after the beginning of fructification, the ecophysiology of the crop, and interactions with pollinating insects (Venturieri, 2011). ...
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In fruit tree breeding, selection indices are used to identify the genotypes that combine desirable commercial and non-commercial characteristics. As Theobroma grandiflorum is generally cultivated in agroforestry systems (AFS), there is a need to develop cultivars that are adapted to such environments. In this study, the objective was to select the most promising genotypes for their future use in AFS based on the additive index, a pioneering method for this crop. The trial was carried out for 12 years in an agroforestry system in the municipality of Tomé-Açu, Pará State, Brazil. The 16 evaluated clones were completely randomised with a variable number of repetitions. The average number of fruits produced as well as the morpho-agronomic characteristics of the fruits were analysed. Mixed linear models were used to estimate the components of variance and predict the genotypic values. The genetic correlation between the variables was estimated, and the selection of genotypes was based on the additive index, with a positive orientation of all variables except the thickness of the fruit shells and the weight of the fruits. Clones 42, 44, 46, 47, 57, 61, and 64 performed well for all the analysed variables, resulting in a selection gain of 7.3% and low incidence rates of witches’ broom disease. These genotypes can be made available to producers in the form of clones for use in AFS and can further be included in future hybridisations in T. grandiflorum breeding.
... Fertilization (allogamic or heterogamic) causes an increase in the ovary which becomes a fruit called «cherelle» (during the growth phase) then «pod» (at the final stage). This maturation varies depending upon the variety and lasts about 4-7 months for cocoa [29,31], and 3-6 months for cupuassu [32]. Both species face fertility challenges, resulting in low fructification rates. ...
... Both species face fertility challenges, resulting in low fructification rates. The known reasons may be (1) lower rates of pollinated flowers, (2) lower effectiveness of entomophilic pollination, and (3) incompatibility reactions inducing flower abortion [29,32]. ...
... Indeed, Cuatrecasas et coll., pointed out that the problem of incompatibility is one of the specificities of the Theobroma species [2]. Cupuassu and cocoa have both been identified as self-incompatible species [32]. Self-incompatibility and cross-incompatibility are defined by the inability to pollinate the flower (by its own pollen or by the pollen from another incompatible tree) to turn it into fruits. ...
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Cocoa and cupuassu are evergreen Amazonian trees belonging to the genus Theobroma, with morphologically distinct fruits, including pods and beans. These beans are generally used for agri-food and cosmetics and have high fat and carbohydrates contents. The beans also contain interesting bioactive compounds, among which are polyphenols and methylxanthines thought to be responsible for various health benefits such as protective abilities against cardiovascular and neurodegenerative disorders and other metabolic disorders such as obesity and diabetes. Although these pods represent 50–80% of the whole fruit and provide a rich source of proteins, they are regularly eliminated during the cocoa and cupuassu transformation process. The purpose of this work is to provide an overview of recent research on cocoa and cupuassu pods and beans, with emphasis on their chemical composition, bioavailability, and pharmacological properties. According to the literature, pods and beans from cocoa and cupuassu are promising ecological and healthy resources.
... Pollinator efficiency is also critical. In T. grandiflorum, it has been found that two-thirds of the flowers remain unpollinated, which indicates that there is little natural pollination (Venturieri 2011). In T cacao, this phenomenon has also been observed in Australia where various strategies have been used to improve polinator's efficiency. ...
... We calculated the percentage of trees with flowers and/ or fruits and graphed the behavior by month following Venturieri's classification for T. grandiflorum (Venturieri 2011) into four categories: 0 = no flowers open, 1 = up to 10 flowers open, 2 = 11-50 flowers open, and 3 = more than 50 flowers. We counted the number of fruits in the crown and graphed the fruiting intensity following also four categories: 0 = no fruits in the crown, 1 = one to five fruits, 2 = six to 15 fruits, and 3 = more than 15 fruits. ...
... Although the correlation was not made in other locations, similar observations were made in Iquitos, Peru (Gonzáles andTorres 2010) andin Ibadan, Nigeria (Aikpokpodion 2012). Similarly, T. grandiflorum was most productive in the driest and most sunny months in Pará, Brazil (Venturieri 2011). This increase in flower productivity is very common in many plants with fragrant flowers to spread the aroma and attract tiny flying pollinators that need to go from one plant to another. ...
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Theobroma bicolor is widely distributed in the Neotropics with great potential for economic use. However, very little is available in literature about its pollinators and reproductive ecology in comparison to what is available about its economically more important relative T. cacao. We studied aspects of the floral biology, reproductive phenology and biotic restrictions to the reproduction of the species. We measured, observed and monitored flowers and described their morphology, breeding system, receptivity times of the stigma and peaks of flower and fruit production. During 20 consecutive days, we collected floral visitors of 135 ± 45 flowers per day. We also calculated the damage in fruit production generated by the mistletoe (Oryctanthus cf. alveolatus) and the fungus (Moniliophthora roreri). We found that flower stigma was more receptive between 6:00-10:00 and flowers last up to three days on the tree, which is less time than what has been reported elsewhere. We collected 211 insects from 68 different morphotypes, but five of these represented 49.8% of the total; specimens of Ceratopogonidae, Chironimidae and Sciaridae were the most abundant and could be pollinators of the species. The flowering of T. bicolor showed a positive correlation (r > 0.75) with the months with higher temperature and solar radiation. We recorded an average per tree of 7 ± 5 fruits. We found a drastic loss of 84% of fruits in the plants infested by the mistletoe O. cf. alveolatus and of 29% when infested by the fungus M. roreri. The results of this study serve as a basis to generate local management practices.
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... The most effective control of witches' broom requires periodic inspections of the plantations for the withdrawal of brooms and dry greens and fruits affected by the disease. This waste must be burned or buried outside the area of planting (Venturieri et al., 1993, Venturieri, 2011 (Figure 1C). None of the 60 farmers studied makes this sanitary control on their cupuassu plantations. ...
... The indiscriminate use of pesticides can lead to problems with pollinators, stressing the fact that no isolated technique to control this pest is available. Pollination deficiency is a limiting factor that affect cupuassu fruit production, as pointed out by Venturieri (2011). This author verified that about 67% of the flowers produced by cupuassu remain unpollinated. ...
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... Although the technique is rarely used for fruit trees and there is no record of its use for cupuassu tree, it is a promising tool that can be employed in genetic improvement. Because cupuassu tree is allogamous and selfincompatible (Venturieri, 2011), it is extremely dependent on genetic variability. It is also a perennial species with a long life cycle. ...
... The Additive Index (AI) enabled us to identify the most promising progeny and, within them, the most appropriate individuals to be cloned, aiming at the exploitation of heterosis resulting from three-way crosses (Prazeres et al., p. 89 2016). The goal was to maintain variability between genotypes by selecting a maximum of two individuals from each progeny, given the self-incompatible, allogamous characteristics of the species, which requires diversity to enable effective crossbreeding (Venturieri, 2011). ...
... The genus 'Theobroma' contains 22 species among which Theobroma cacao L. is widely cultivated. Theobroma grandiflorum L. is the other closely related species of cocoa, which is also the source for a variety of chocolate known as cupulate or cupuacu (Venturieri, 2011). The tropical plant is a native of Amazon region of South America (Bartley, 2005;Cheesman, 1944); later its cultivation spread to the countries in Asia and Africa (Bartley, 2005;Zhang and Motilal, 2016). ...
... Though, a small number of pollinated flowers developed into fruits is a common physiological characteristic of young [24][25][26]. Venturieri [27] reported that for crop species with fruits which are energetically expensive to produce such as cocoa, only few fruits will be produced. This is because the mother tree is physiologically unable to provide all the nutrient requirements for both flower and fruit growths [28]. ...
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... This highlights the impact the type of management can have on the phenotypic manifestation of the evaluated traits, especially mean fruit production/plant. This fact, combined with the variability resulting from the species' self-incompatibility (Venturieri 2011), is reflected in uncertainties about the cultivation of genetic materials that have not been evaluated in a range of environments and emphasizes the importance of studies of this nature. ...
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The cultivation of Theobroma grandiflorum in the Brazilian Amazon is mainly conducted by family farmers who use a range of different management straegies. Thus, breeding programs of the species must address the challenge of developing cultivars that are adapted to and stable in a variety of cultivation environments. In this context, this study aimed to estimate the optimum number of harvests for genetic selection of T. grandiflorum progenies and identify the most promising ones in terms of productivity, stability, and adaptability. The trials were implemented in three environments, using a randomized complete block design, with 25 full-sib progenies, five replications, and three plants per plot. The traits mean number of fruits/plant, mean fruit production/plant, and rate of infection with witches’ broom ( Moniliophthora perniciosa ) were evaluated over 11 harvests. The Restricted Maximum Likelihood/Best Linear Unbiased Prediction (REML/BLUP) mixed model method was used to estimate genetic parameters and predict genetic values, which were then applied to assess stability and adaptability. The results show that there is genetic variability among the studied T. grandiflorum progenies and that accurate genetic selection aiming at recombination is effective after three harvests, for recombination, or eleven harvests for identification of recommended progenies. Six progenies were selected that met the requirements for productivity, stability, and adaptability to different cultivation environments. These results can be used to optimize and advance T. grandiflorum breeding programs.
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In the present work, cacauhy's (Theobroma speciosum) floral biology was studied. Flower buds split their sepals at 14h reaching its maximum at 22h, but all flowers were fully opened at 6:00 h of the following morning. Stigmatic branches showed exudates, reaching maximum between 6:00 h and 10:00 h at the same day. Ligules and petal hoods were the floral parts with highest intensity of odour. Flowers were receptive along all the morning and noon of the anthesis day. Approximately 65% of the flowers were naturally pollinated, but only 0.85% of them set a fruit. Abscission occurred on its higher frequency at 6:00 h of the second day after anthesis. Controlled pollinations showed that cacauhy was self-incompatible species.
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The premier text in the field, Biometry provides both an elementary introduction to basic biostatistics as well as coverage of more advanced methods used in biological research. Students are shown how to think through research problems and understand the logic behind the different experimental situations. This book is designed to serve not only as a text to accompany a lecture course but is also a must-have reference text! NEW TO THIS EDITION • An Increased Focus on Computer-Based Statistical Methods. Computational formulas have also been replaced throughout with simpler structural formulas for ease of understanding. • Matrix methods. Matrix methods are introduced in new sections on multiple regression, general linear models, ancova, and curvilinear regression. A new appendix on matrix algebra is also included. • New Chapter on Statistical Power and Sample Size Estimation. The new edition features a new chapter that covers statistical power, measures of effect size, and the estimation of sample size required for tests and for confidence limits. • Up-to-Date Coverage of Key Developments in Biostatistics. This edition includes the most up-to-date coverage of key topics such as meta-analysis and trends in the discipline such as the use of resampling methods.
Article
The evolution of selfing vs. outbreeding has been of major interest to plant population biology. The independent historic introductions of self-compatible and self-incompatible genotypes of cacao in Trinidad have allowed us to study the selection acting upon an unnatural breeding system polymorphism. Field observations of an abandoned cacao plantation indicate that the self-incompatible phenotype has slightly increased in frequency within a single generation. The self-compatible trees produced significantly less flowers but still set more pods than did the self-incompatible trees, although compatibility types did not differ in tree size or mature seed production. Greenhouse observations suggest that the apparent failure of self-compatibility to increase in the population is related to inbreeding depression resulting from selfing, expressed as reduced seedling establishment.