Cocoa Breeding and Genetic Resources Utilisation

Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

[68] Nortey TN, Boadu S, Naazie A. Effect of phytase supplementation on performance of grower pigs fed diets based on maize, wheat bran and reducing levels of soybean meal, but low in nutrient density. Ghanaian Journal of Animal Science. 2013;**7**(2):117-122

[69] Nortey TN, Patience JF,

[70] Alemawor F, Oddoye EOK, Dzogbefia VP, Oldham JH, Donkor A. Broiler performance on finisher diets containing different levels of either *Pleurotus ostreatus*-fermented dried cocoa pod husk or dried cocoa pod husk supplemented with enzymes. Tropical Animal Health and Production.

[71] Nortey TN, Kpogo DV, Kpogo AL, Naazie A, Oddoye EOK. Cocoa pod husk is a potential feed ingredient in laying hen diets. Livestock Research for Rural Development. 2015;**27**(6):200-205

2010;**42**:933-939

Simmins PH, Trotier NL, Zijlstra RT. Effects of individual or combined xylanase and phytase supplementation on energy, amino acid, and phosphorus digestibility and growth performance of grower pigs fed wheat-based diets containing wheat millrun. Journal of Animal Science. 2007;**85**:1432-1443

**100**

**103**

**Chapter 6**

**Abstract**

industry in the country are discussed.

**1. Introduction**

Cacao Genetic Resources

Conservation and Utilization for

Sustainable Production in Nigeria

*Theobroma cacao*, the source of chocolate, is one of the most important tree-crop that serves the purpose of sustaining the economy of millions of households and the largest non-oil foreign exchange earnings in Nigeria. The management of cacao genetic resources as it affects sustainable production of cocoa in Nigeria is reviewed. These include details of the diversity present in the germplasm collections, their utilization in varietal development and current status of the field genebanks as revealed by DNA fingerprinting using single nucleotide polymorphisms (SNPs) markers. Factors contributing to continuing backlash in the annual outputs of cocoa in Nigeria are also highlighted. The prospects of advances in the science of cacao genomics for up-scaling production and its impacts on the improvement of the

*Festus Olakunle Olasupo and Peter O. Aikpokpodion*

**Keywords:** germplasm introduction, genetic diversity, cacao breeding,

*Theobroma cacao* L., a member of Malvaceae family [1] is a small under-story tree that produces cocoa beans used in the manufacture of chocolate, cosmetics, confectioneries and other cocoa products. Recent evident from restricted fragment length polymorphisms (RFLP) and microsatellite analyses of ancient Criollo trees, consolidated proposition of the humid tropics South American Amazon regions as its center of origin [2]. Cacao is now grown as a tree crop in the tropical regions of the world between latitudes 20° north and 20° south of the equator [3]. Cocoa is one of the most economically important agricultural commodities in Africa and has been contributing to gross domestic product (GDP), national income (NI) and foreign exchange earnings of many African producing nations. It is marketed in The United States of America, Europe, United Kingdom and Asia where its butter and solids are used by processing industries. The global exports value of dried beans is between USD 8–10 billion per annum and there has been increasing demand for chocolate in the developing economies of Brazil, China, Eastern Europe and India [4]. In West and Central Africa, more than 96% of cocoa production is carried out by smallholder farmers who rely on proceeds from cocoa beans as a major source of family income [5, 6]. The world produced 4.744 million metric tons of dried beans in 2017 and 76% of the world production comes from Africa with about 7% of the continent exports from Nigeria, ranking the country 6th in the world [4]. In Nigeria,

cacao genomics, cacao science, sustainable production

## **Chapter 6**

## Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria

*Festus Olakunle Olasupo and Peter O. Aikpokpodion*

## **Abstract**

*Theobroma cacao*, the source of chocolate, is one of the most important tree-crop that serves the purpose of sustaining the economy of millions of households and the largest non-oil foreign exchange earnings in Nigeria. The management of cacao genetic resources as it affects sustainable production of cocoa in Nigeria is reviewed. These include details of the diversity present in the germplasm collections, their utilization in varietal development and current status of the field genebanks as revealed by DNA fingerprinting using single nucleotide polymorphisms (SNPs) markers. Factors contributing to continuing backlash in the annual outputs of cocoa in Nigeria are also highlighted. The prospects of advances in the science of cacao genomics for up-scaling production and its impacts on the improvement of the industry in the country are discussed.

**Keywords:** germplasm introduction, genetic diversity, cacao breeding, cacao genomics, cacao science, sustainable production

## **1. Introduction**

*Theobroma cacao* L., a member of Malvaceae family [1] is a small under-story tree that produces cocoa beans used in the manufacture of chocolate, cosmetics, confectioneries and other cocoa products. Recent evident from restricted fragment length polymorphisms (RFLP) and microsatellite analyses of ancient Criollo trees, consolidated proposition of the humid tropics South American Amazon regions as its center of origin [2]. Cacao is now grown as a tree crop in the tropical regions of the world between latitudes 20° north and 20° south of the equator [3]. Cocoa is one of the most economically important agricultural commodities in Africa and has been contributing to gross domestic product (GDP), national income (NI) and foreign exchange earnings of many African producing nations. It is marketed in The United States of America, Europe, United Kingdom and Asia where its butter and solids are used by processing industries. The global exports value of dried beans is between USD 8–10 billion per annum and there has been increasing demand for chocolate in the developing economies of Brazil, China, Eastern Europe and India [4]. In West and Central Africa, more than 96% of cocoa production is carried out by smallholder farmers who rely on proceeds from cocoa beans as a major source of family income [5, 6]. The world produced 4.744 million metric tons of dried beans in 2017 and 76% of the world production comes from Africa with about 7% of the continent exports from Nigeria, ranking the country 6th in the world [4]. In Nigeria, cocoa provides means of livelihood to more than 5 million people and serves the purpose of sustaining the economy of millions of households to live above poverty and hunger. Although most of the country's budgetary revenue comes from sale of crude oil, agriculture contributes significantly to the economy with about 70% of the population engaged in agriculture. Income obtained from cocoa exports accounts for up to 27% of the 41.48% of Nigeria's GDP attributed to agriculture [7] and it is the largest non-oil foreign exchange earnings. However, cocoa production in Nigeria has not been sustainable as it reflected in the decline of the country's export data observed for almost two decades. Many factors are responsible for the low farm productivity which in turn is contributing to continuing instability and negative downturn in cocoa production in Nigeria [8]. These include but not limited to climate variability, diseases and pests infestations and poor access to improved planting materials. Development of improved hybrid varieties with good yield quality potentials is a necessity for a sustainable cocoa production system and this is subject to the amount of genetic variability available within germplasm collection. Thus, determined and consistent efforts are required in the acquisition of the needed genetic materials/resources for their conservation, evaluation and efficient use to develop well-adapted cultivars for combating the cocoa production challenges.

## **2. Germplasm introduction**

Cacao was first introduced into Africa from Bahia, the Amazonian Region in Brazil by the Portuguese to Principe in 1822, with the establishment of 30 cacao plants of the Amelonado type (Lower Amazon Forastero) in Principe and later expanded to the neighboring island of Sao Tome [3]. Toxopeus [9] also reported that the two cacao trees of the Amelonado variety which were successfully established in Sao Tome from a batch of plants brought in from Bahia, Brazil in 1822, became the parents of the subsequent cacao trees of the island. Further movement of cacao from Sao Tome Principe islands to Fernando Po (the present-day Bioko in Equatorial Guinea) took place in 1855 by the Spanish seafarers [9, 10]. Other introductions were also made by Swiss missionaries from Suriname, and the first set of cocoa seeds were sown on Africa mainland in 1857 [11]. However, the variety which was originally brought into the island from Brazil was in its adopted home known as 'Sao Tome Creolou,' to distinguish it from other varieties that had been introduced subsequently. Following the introduction of the variety into West Africa and its wide cultivation in the region, it became popularly referred to as 'West African Amelonado.'

The history of cacao genetic materials introduction into Africa and subsequently into Nigeria is simply illustrated in (**Figure 1**). Cacao germplasm introduction to Nigeria can be presented as a sub-set of its introduction to Africa using the two phases partitioning ideology (Exploratory Colonial Period and Expansionary Experimental Pre- and Post-Independence Period) as reviewed by Aikpokpodion [7].

#### **2.1 Exploratory colonial period (1874–1909)**

The first era of cacao introduction into Nigeria started in 1874 by Chief Squiss Ibaningo, a trader who transported pods of Amelonado cocoa from Fernando Po into Bonny (now Rivers State of Nigeria) [1]. Many of the liberated slaves who were settled on the Fernando Po Island who had worked as contract labourers in the local cocoa plantations also contributed to cacao introduction into Nigeria. In 1880, there was an evidence of cacao plantation near Agege, owned by J.P.L Davies, most probably a

**105**

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria*

liberated slave [13], who had been the captain of a freighter plying the West African coast and could have obtained the seed from Fernando Po. The first cacao seedlings were planted somewhere near Ibadan prior to 1890 [14]. Berry [15] also reported that Ogunwale was the first to carry about 200 cacao pods to Ibadan for planting from Agege, which must have been from Davies' plantation. Furthermore, around 1900, the Colonial administration also introduced some red-podded cacao (Trinitario) materials to the botanical garden established in Agege Lagos in 1888 from British West Indies, presumably by the Jamaican curator of the garden. Several cacao germplasm materials were also introduced into Nigeria by the missionaries and slave traders. As a result of these various introduction efforts, a fair range of variability was presumably present on farmers' plantations with self-compatible 'West African Amelonado' types dominating the complex mixture of cacao of diverse origin. Towards the end of nineteenth century, cocoa cultivation and industry had started with 95% of national

*Points of cacao introduction into Africa (Sao Tome and Principe) and Nigeria (Bonny Island). Place of* 

At the beginning of twentieth century, cocoa was already a valuable commodity crop in Nigeria. Therefore, further introductions of cacao genetic materials during the second era were for economic reasons with the aims of obtaining better income and premium due to greater yields and higher bean and chocolate quality. However, during the last decade, the "People, Planet and Profit" concept of sustainability has become a significant factor in cacao germplasm introduction. This has bearing with the concept of "Preventive Breeding" where clones showing resistance to regionally important diseases of cocoa growing regions could be introduced through international intermediate quarantine centers [7]. This was to ensure that there is present in the germplasm collections, adequate sources of resistance to cope with, in case of new disease spread in order to prevent local cocoa economy crash that is associated

Series of additional germplasm introductions following the initial successful introduction of the Brazilian Amelonado of the Lower Amazon Forastero type have

export of 21 tons in 1895 coming from the Western Region [12].

*first introduction to Africa in 1822. Place of first introduction to Nigeria in 1857.*

**2.2 Expansionary experimental pre- and post-independence** 

**period (1910–2018)**

**Figure 1.**

with any epidemics.

*DOI: http://dx.doi.org/10.5772/intechopen.82703*

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria DOI: http://dx.doi.org/10.5772/intechopen.82703*

#### **Figure 1.**

Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

cocoa provides means of livelihood to more than 5 million people and serves the purpose of sustaining the economy of millions of households to live above poverty and hunger. Although most of the country's budgetary revenue comes from sale of crude oil, agriculture contributes significantly to the economy with about 70% of the population engaged in agriculture. Income obtained from cocoa exports accounts for up to 27% of the 41.48% of Nigeria's GDP attributed to agriculture [7] and it is the largest non-oil foreign exchange earnings. However, cocoa production in Nigeria has not been sustainable as it reflected in the decline of the country's export data observed for almost two decades. Many factors are responsible for the low farm productivity which in turn is contributing to continuing instability and negative downturn in cocoa production in Nigeria [8]. These include but not limited to climate variability, diseases and pests infestations and poor access to improved planting materials. Development of improved hybrid varieties with good yield quality potentials is a necessity for a sustainable cocoa production system and this is subject to the amount of genetic variability available within germplasm collection. Thus, determined and consistent efforts are required in the acquisition of the needed genetic materials/resources for their conservation, evaluation and efficient use to develop well-adapted cultivars for combating the cocoa production

Cacao was first introduced into Africa from Bahia, the Amazonian Region in Brazil by the Portuguese to Principe in 1822, with the establishment of 30 cacao plants of the Amelonado type (Lower Amazon Forastero) in Principe and later expanded to the neighboring island of Sao Tome [3]. Toxopeus [9] also reported that the two cacao trees of the Amelonado variety which were successfully established in Sao Tome from a batch of plants brought in from Bahia, Brazil in 1822, became the parents of the subsequent cacao trees of the island. Further movement of cacao from Sao Tome Principe islands to Fernando Po (the present-day Bioko in Equatorial Guinea) took place in 1855 by the Spanish seafarers [9, 10]. Other introductions were also made by Swiss missionaries from Suriname, and the first set of cocoa seeds were sown on Africa mainland in 1857 [11]. However, the variety which was originally brought into the island from Brazil was in its adopted home known as 'Sao Tome Creolou,' to distinguish it from other varieties that had been introduced subsequently. Following the introduction of the variety into West Africa and its wide cultivation in the region, it became popularly referred to as 'West

The history of cacao genetic materials introduction into Africa and subsequently into Nigeria is simply illustrated in (**Figure 1**). Cacao germplasm introduction to Nigeria can be presented as a sub-set of its introduction to Africa using the two phases partitioning ideology (Exploratory Colonial Period and Expansionary

Experimental Pre- and Post-Independence Period) as reviewed by Aikpokpodion [7].

The first era of cacao introduction into Nigeria started in 1874 by Chief Squiss Ibaningo, a trader who transported pods of Amelonado cocoa from Fernando Po into Bonny (now Rivers State of Nigeria) [1]. Many of the liberated slaves who were settled on the Fernando Po Island who had worked as contract labourers in the local cocoa plantations also contributed to cacao introduction into Nigeria. In 1880, there was an evidence of cacao plantation near Agege, owned by J.P.L Davies, most probably a

**104**

challenges.

**2. Germplasm introduction**

African Amelonado.'

**2.1 Exploratory colonial period (1874–1909)**

*Points of cacao introduction into Africa (Sao Tome and Principe) and Nigeria (Bonny Island). Place of first introduction to Africa in 1822. Place of first introduction to Nigeria in 1857.*

liberated slave [13], who had been the captain of a freighter plying the West African coast and could have obtained the seed from Fernando Po. The first cacao seedlings were planted somewhere near Ibadan prior to 1890 [14]. Berry [15] also reported that Ogunwale was the first to carry about 200 cacao pods to Ibadan for planting from Agege, which must have been from Davies' plantation. Furthermore, around 1900, the Colonial administration also introduced some red-podded cacao (Trinitario) materials to the botanical garden established in Agege Lagos in 1888 from British West Indies, presumably by the Jamaican curator of the garden. Several cacao germplasm materials were also introduced into Nigeria by the missionaries and slave traders. As a result of these various introduction efforts, a fair range of variability was presumably present on farmers' plantations with self-compatible 'West African Amelonado' types dominating the complex mixture of cacao of diverse origin. Towards the end of nineteenth century, cocoa cultivation and industry had started with 95% of national export of 21 tons in 1895 coming from the Western Region [12].

## **2.2 Expansionary experimental pre- and post-independence period (1910–2018)**

At the beginning of twentieth century, cocoa was already a valuable commodity crop in Nigeria. Therefore, further introductions of cacao genetic materials during the second era were for economic reasons with the aims of obtaining better income and premium due to greater yields and higher bean and chocolate quality. However, during the last decade, the "People, Planet and Profit" concept of sustainability has become a significant factor in cacao germplasm introduction. This has bearing with the concept of "Preventive Breeding" where clones showing resistance to regionally important diseases of cocoa growing regions could be introduced through international intermediate quarantine centers [7]. This was to ensure that there is present in the germplasm collections, adequate sources of resistance to cope with, in case of new disease spread in order to prevent local cocoa economy crash that is associated with any epidemics.

Series of additional germplasm introductions following the initial successful introduction of the Brazilian Amelonado of the Lower Amazon Forastero type have been reviewed by authors [3, 7, 16, 17]. The 'Trinitario' and 'Criollo' types were introduced to Nigeria in 1920 which formed hybrids with the original Amelonado type [18]. The former Nigeria Department of Agriculture was established in 1910 at Moor plantation, Ibadan. Nevertheless, the formal germplasm conservation and selection programs started around 1931 by O.J. Voelckler at the Nigerian Department of Agriculture in Moor Plantation, Ibadan under the Colonial Administration [17] with the sole aim of development and release of improved planting materials. In 1933, further germplasm introductions of some Trinitario and Criollo selections were made respectively, from Trinidad and Ceylon (the present day Sri Lanka) [19]. However, the outbreak of cocoa swollen shoot virus disease in the 1930s in Ghana, Togo and Nigeria almost destroyed the cocoa industry due to narrow genetic base in the population. This led to the establishment of West Africa Cocoa Research Institute (WACRI) in 1944 with the headquarters in Tafo, Ghana and a sub-station in Ibadan by an inter-territorial research thrust under the British West African Colonial Administration. Following the institute's establishment, several accessions from Upper Amazon Forastero and 'Trinitario' populations collected by Pound [20, 21] in Trinidad were introduced in 1944 into Tafo in Ghana by WACRI to widen the genetic base of the germplasm [22, 23] and subsequently into Nigeria from Ghana. Open pollinated pods from 11 selected accessions of the Upper Amazon Trinidad introduction that were established in Ghana were also introduced into Ibadan, Nigeria to form the F2 Amazon population and these form the source of the open pollinated "F3 Amazon" or "Mixed Amazon" materials in Nigeria up till present time.

The Nigeria sub-station of WACRI was upgraded to national research station, the Cocoa Research Institute of Nigeria (CRIN) in 1964 to focus on research that will facilitate improved production of cocoa. Then cacao seeds from 10 different crosses were introduced from Wageningen (The Netherlands) between 1964 and 1965 to raise 390 hybrid seedlings established in CRIN. Between 1965 and 1967, a large-scale introduction of Upper Amazon cacao materials was made from Trinidad as part of the Trinidad-Nigeria Cacao Introduction Scheme sponsored by the Cocoa Alliance [24]. This consisted of 313 clones and 701 seedling progenies of intra-Nanay, intra-Parinari, intra-Iquitos and inter-P (Pound's selections) crosses derived from a total of 350 crosses [25]. These clones and hybrids introduced from Trinidad constituted the "T clones" of CRIN germplasm collections. Materials were also acquired from Costa Rica, Indonesia, Fernando Po, Kew Gardens (United Kingdom) and Miami (USA) [19]. Between 1998 and 2004, 43 clones were introduced into Nigeria through an international initiative known as "Cocoa Germplasm Utilization and Utilization: A Global Approach," a project sponsored by the United Nations Common Fund for Commodity (CFC), through the supervision of the International Plant Genetic Resources Institute (IPGRI) as the executing agency [26]. Many other genetic materials introductions have not been reported adequately purposely because it is considered of hardly any use to publish introductions made if most of them did not survive due to the difficulties in their establishment. In the late 2017 and early 2018, some clones were introduced from International Cocoa Quarantine Center, University of Reading, of which 32 clones were established in a new germplasm plot at CRIN, Ibadan.

## **3. Status of Nigeria cacao field genebank**

The genetic resources of cacao are composed of genetic variability that serve as the raw materials for the development of new varieties and cultivar improvement to enhance the production system for sustaining cocoa industry. These materials are conserved *ex situ* in field genebanks (germplasm plots of CRIN) and various

**107**

**Figure 2.**

*Population structure indicating cacao types grown on farmers' fields in Nigeria.*

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria*

seed garden plots in major producing state of the country as well as *in situ*, that is in farmers' fields across the growing ecology. Appreciable diversity among these materials is of fundamental importance in the sustainability of cocoa production as it provides the necessary adaptation to the prevailing biotic and abiotic environmental challenges and enables changes in the genetic composition to cope with changes in the environment. The set of materials introduced during the Colonial era and the widely cultivated cacao in Nigeria in the early twentieth century (Amelonado and Trinitario varieties) were known to have narrow genetic base. However, the incidence of CSSV epidemics in the 1930s led to the introduction of Upper Amazon materials into the country's cacao genepool. Furthermore, several targeted germplasm collection efforts were made from mid-1960s up till recent times by CRIN breeding programme to broaden the diversity of Nigeria cacao genetic base. Research reports from the studies of genebanks collections and cacao accessions collected from farmers' fields in growing ecologies using 12 microsatellite markers showed that there is an appreciable improvement in Nigeria cacao genetic diversity [7, 16, 17, 27]. Of the 574 accessions studied, a total of 144 alleles were detected with a mean allelic richness of 4.39 alleles per locus. Upper Amazon parent population was observed to have largest genetic diversity (Hnb = 0.730), followed by the Posnette's Introduction (Hnb = 0.704) while the least (Hnb = 0.471) was recorded in the local parent population. Population structure analysis of cacao types grown in farmer's fields showed that the Upper Amazon Forastero, Amelonado, Trinitario and others constitute 66, 24, 6 and 4% of cacao grown, respectively (**Figure 2**). More recent diversity studies using simple nucleotide polymorphism (SNP) markers [28], revealed the presence of cacao genetic groups that cut across the major primary populations in Nigeria germplasm collection (**Figure 3**) thus indicating appreciable improvement in the genetic base. This progress has been attributed to the international clones recently introduced into the country's cacao collection. However, there is paucity of report on the evaluation of recently introduce germplasm materials to enhance their utility in varietal development. This may have been responsible for authors' reports that a small proportion of the genetic diversity available in field genebanks at CRIN had been used to develop improved varieties supplied to farmers as planting materials [7, 17, 28]. In addition to this, the impact of mislabeling and off-type among the parental clones in the seed gardens and germplasm accessions of Nigerian field genebanks have been

*DOI: http://dx.doi.org/10.5772/intechopen.82703*

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria DOI: http://dx.doi.org/10.5772/intechopen.82703*

seed garden plots in major producing state of the country as well as *in situ*, that is in farmers' fields across the growing ecology. Appreciable diversity among these materials is of fundamental importance in the sustainability of cocoa production as it provides the necessary adaptation to the prevailing biotic and abiotic environmental challenges and enables changes in the genetic composition to cope with changes in the environment. The set of materials introduced during the Colonial era and the widely cultivated cacao in Nigeria in the early twentieth century (Amelonado and Trinitario varieties) were known to have narrow genetic base. However, the incidence of CSSV epidemics in the 1930s led to the introduction of Upper Amazon materials into the country's cacao genepool. Furthermore, several targeted germplasm collection efforts were made from mid-1960s up till recent times by CRIN breeding programme to broaden the diversity of Nigeria cacao genetic base. Research reports from the studies of genebanks collections and cacao accessions collected from farmers' fields in growing ecologies using 12 microsatellite markers showed that there is an appreciable improvement in Nigeria cacao genetic diversity [7, 16, 17, 27]. Of the 574 accessions studied, a total of 144 alleles were detected with a mean allelic richness of 4.39 alleles per locus. Upper Amazon parent population was observed to have largest genetic diversity (Hnb = 0.730), followed by the Posnette's Introduction (Hnb = 0.704) while the least (Hnb = 0.471) was recorded in the local parent population. Population structure analysis of cacao types grown in farmer's fields showed that the Upper Amazon Forastero, Amelonado, Trinitario and others constitute 66, 24, 6 and 4% of cacao grown, respectively (**Figure 2**).

More recent diversity studies using simple nucleotide polymorphism (SNP) markers [28], revealed the presence of cacao genetic groups that cut across the major primary populations in Nigeria germplasm collection (**Figure 3**) thus indicating appreciable improvement in the genetic base. This progress has been attributed to the international clones recently introduced into the country's cacao collection. However, there is paucity of report on the evaluation of recently introduce germplasm materials to enhance their utility in varietal development. This may have been responsible for authors' reports that a small proportion of the genetic diversity available in field genebanks at CRIN had been used to develop improved varieties supplied to farmers as planting materials [7, 17, 28]. In addition to this, the impact of mislabeling and off-type among the parental clones in the seed gardens and germplasm accessions of Nigerian field genebanks have been

**Figure 2.** *Population structure indicating cacao types grown on farmers' fields in Nigeria.*

Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

Amazon" or "Mixed Amazon" materials in Nigeria up till present time.

The Nigeria sub-station of WACRI was upgraded to national research station, the Cocoa Research Institute of Nigeria (CRIN) in 1964 to focus on research that will facilitate improved production of cocoa. Then cacao seeds from 10 different crosses were introduced from Wageningen (The Netherlands) between 1964 and 1965 to raise 390 hybrid seedlings established in CRIN. Between 1965 and 1967, a large-scale introduction of Upper Amazon cacao materials was made from Trinidad as part of the Trinidad-Nigeria Cacao Introduction Scheme sponsored by the Cocoa Alliance [24]. This consisted of 313 clones and 701 seedling progenies of intra-Nanay, intra-Parinari, intra-Iquitos and inter-P (Pound's selections) crosses derived from a total of 350 crosses [25]. These clones and hybrids introduced from Trinidad constituted the "T clones" of CRIN germplasm collections. Materials were also acquired from Costa Rica, Indonesia, Fernando Po, Kew Gardens (United Kingdom) and Miami (USA) [19]. Between 1998 and 2004, 43 clones were introduced into Nigeria through an international initiative known as "Cocoa Germplasm Utilization and Utilization: A Global Approach," a project sponsored by the United Nations Common Fund for Commodity (CFC), through the supervision of the International Plant Genetic Resources Institute (IPGRI) as the executing agency [26]. Many other genetic materials introductions have not been reported adequately purposely because it is considered of hardly any use to publish introductions made if most of them did not survive due to the difficulties in their establishment. In the late 2017 and early 2018, some clones were introduced from International Cocoa Quarantine Center, University of Reading, of which 32 clones were established in a

The genetic resources of cacao are composed of genetic variability that serve as the raw materials for the development of new varieties and cultivar improvement to enhance the production system for sustaining cocoa industry. These materials are conserved *ex situ* in field genebanks (germplasm plots of CRIN) and various

been reviewed by authors [3, 7, 16, 17]. The 'Trinitario' and 'Criollo' types were introduced to Nigeria in 1920 which formed hybrids with the original Amelonado type [18]. The former Nigeria Department of Agriculture was established in 1910 at Moor plantation, Ibadan. Nevertheless, the formal germplasm conservation and selection programs started around 1931 by O.J. Voelckler at the Nigerian Department of Agriculture in Moor Plantation, Ibadan under the Colonial Administration [17] with the sole aim of development and release of improved planting materials. In 1933, further germplasm introductions of some Trinitario and Criollo selections were made respectively, from Trinidad and Ceylon (the present day Sri Lanka) [19]. However, the outbreak of cocoa swollen shoot virus disease in the 1930s in Ghana, Togo and Nigeria almost destroyed the cocoa industry due to narrow genetic base in the population. This led to the establishment of West Africa Cocoa Research Institute (WACRI) in 1944 with the headquarters in Tafo, Ghana and a sub-station in Ibadan by an inter-territorial research thrust under the British West African Colonial Administration. Following the institute's establishment, several accessions from Upper Amazon Forastero and 'Trinitario' populations collected by Pound [20, 21] in Trinidad were introduced in 1944 into Tafo in Ghana by WACRI to widen the genetic base of the germplasm [22, 23] and subsequently into Nigeria from Ghana. Open pollinated pods from 11 selected accessions of the Upper Amazon Trinidad introduction that were established in Ghana were also introduced into Ibadan, Nigeria to form the F2 Amazon population and these form the source of the open pollinated "F3

**106**

new germplasm plot at CRIN, Ibadan.

**3. Status of Nigeria cacao field genebank**

#### **Figure 3.**

*Population structure of Nigeria cacao germplasm collections.*


#### **Table 1.**

*Examples of DNA fingerprints based on multi-locus matching of 28 SNPs between original references and Nigerian cacao collection (showing truncated profiles).*

a significant problem hindering their efficient conservation and use for breeding programs. The occurrence of mislabeling and offtypes was first observed in the Nigerian cocoa germplasm collection by the application of SSR markers for genotyping [17]. This was recently consolidated by Olasupo et al. [28] through the application of SNPs for DNA fingerprinting of field genebank collections in which high level of mislabeling was detected in the recently introduced international germplasm materials (**Table 1**). Mislabeling has been identified as one of the key factors contributing to the high rate of unwanted/unproductive progenies produced in seed gardens because this can seriously compromise the quality of seedlings that would be distributed to farmers [17, 29]. Labeling errors in cacao field genebanks have been attributed to error from the source of germplasm introduction [30], human mislabeling errors in the nursery [31], sprouted rootstocks overtaken the scions due to poor germplasm management and multiplied error effect of using wrongly identified clones for new seed garden establishment [28].

In addition, the genetic diversity of cacao that constitute valuable resources in field genebanks useful to sustain cocoa production in Nigeria are further threatened with the following challenges:

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HH268 and IS36.

menace.

were obtained from this programme.

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria*

1.Lack of funding for long-term management of collections, for research on

2.The field genebank collections are predisposed to the challenge of climate

3.Insufficient well trained personnel (expertise) for effective collection, conser-

4.Urbanization and scarcity of land have led to the loss of many germplasm plots

6.Loss of germplasm to natural disaster (such as fire outbreak). There is the need for duplication of most of the field \*\*\*genebanks to serve as back-up (prefer-

7.Insufficient genetic variation in the collections to enhance effective selection

Therefore, there is the need for all the stakeholders to protect the diversity of cacao germplasm collections with the aims of continuous supply of planting materials that have great potentials for high yield, disease and pest resistance, good

architecture, drought tolerance and excellent flavor quality in the future.

Cacao breeding research in Nigeria can be partitioned into four phase: *First Breeding Phase (1931–1956):* The first phase of Nigeria cacao breeding started in 1931 [19]. Some of the trials conducted include the progeny trials of 1942 and 1945 by Voelcker which was the first of its kind in cacao research worldwide. Other trials conducted within this period include clonal trial in 1953 and the double cross hybrid vigour trial in 1954. The major breeding challenge during this period was low level of genetic variability among the parental accession. The main output of the programme was the identification of some materials with hybrid vigour and clonal propagation of the materials. These were N38 and other clonal selections— NT (Nigerian-Trinidad hybrid) 39, 114, 164, 215, 216, 284, 310, and 655. Others selections from the local clones and West African Amelonado population were

In 1938, the West African Cocoa Research Institute (WACRI) started a *Cocoa Improvement and Varietal Development Programme* with the main objective of dealing with the threat posed by the cocoa swollen shoot virus (CSSV) disease in the West African sub region, particularly Ghana and Nigeria. The two main outputs

i.First, introduction of the Upper Amazon genetic materials into Ghana and Nigeria cacao genepool which was very efficient in combatting the CSSV

ii.Two general purpose varieties were also obtained from the programme. The F3-Amazon—a third generation progenies resulting from open pollination

**4. Breeding efforts and impacts on national output**

5.Problem of old age germplasm plots and the need to rejuvenate the old trees

thereby resulting into extinction of some valuable genetic resources.

variability and its associated diseases and pests outbreak

vation and management of germplasm materials

*DOI: http://dx.doi.org/10.5772/intechopen.82703*

diversity, evaluation and use in breeding

ably *in vitro* or cryo-preservation)

of some specific traits.

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria DOI: http://dx.doi.org/10.5772/intechopen.82703*


Therefore, there is the need for all the stakeholders to protect the diversity of cacao germplasm collections with the aims of continuous supply of planting materials that have great potentials for high yield, disease and pest resistance, good architecture, drought tolerance and excellent flavor quality in the future.

## **4. Breeding efforts and impacts on national output**

Cacao breeding research in Nigeria can be partitioned into four phase: *First Breeding Phase (1931–1956):* The first phase of Nigeria cacao breeding started in 1931 [19]. Some of the trials conducted include the progeny trials of 1942 and 1945 by Voelcker which was the first of its kind in cacao research worldwide. Other trials conducted within this period include clonal trial in 1953 and the double cross hybrid vigour trial in 1954. The major breeding challenge during this period was low level of genetic variability among the parental accession. The main output of the programme was the identification of some materials with hybrid vigour and clonal propagation of the materials. These were N38 and other clonal selections— NT (Nigerian-Trinidad hybrid) 39, 114, 164, 215, 216, 284, 310, and 655. Others selections from the local clones and West African Amelonado population were HH268 and IS36.

In 1938, the West African Cocoa Research Institute (WACRI) started a *Cocoa Improvement and Varietal Development Programme* with the main objective of dealing with the threat posed by the cocoa swollen shoot virus (CSSV) disease in the West African sub region, particularly Ghana and Nigeria. The two main outputs were obtained from this programme.


Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

a significant problem hindering their efficient conservation and use for breeding programs. The occurrence of mislabeling and offtypes was first observed in the Nigerian cocoa germplasm collection by the application of SSR markers for genotyping [17]. This was recently consolidated by Olasupo et al. [28] through the application of SNPs for DNA fingerprinting of field genebank collections in which high level of mislabeling was detected in the recently introduced international germplasm materials (**Table 1**). Mislabeling has been identified as one of the key factors contributing to the high rate of unwanted/unproductive progenies produced in seed gardens because this can seriously compromise the quality of seedlings that would be distributed to farmers [17, 29]. Labeling errors in cacao field genebanks have been attributed to error from the source of germplasm introduction [30], human mislabeling errors in the nursery [31], sprouted rootstocks overtaken the scions due to poor germplasm management and multiplied error effect of using wrongly identified clones for new seed garden establishment [28]. In addition, the genetic diversity of cacao that constitute valuable resources in field genebanks useful to sustain cocoa production in Nigeria are further threatened

*Examples of DNA fingerprints based on multi-locus matching of 28 SNPs between original references and* 

**108**

**Figure 3.**

**Table 1.**

*Population structure of Nigeria cacao germplasm collections.*

*Nigerian cacao collection (showing truncated profiles).*

with the following challenges:

of the eleven approved Amazon T (Trinidad) types having broad parental background and have been shown to be superior to West African Amelonado in establishment ability, vegetative vigor, precocity (earliness in bearing) and yield. F3-Amazon variety is also tolerant of CSSV and capsid attack [23, 32]. The Mixed Series II Hybrids (WACRI Series II) varieties were produced from crosses of Upper Amazon cacao with local selections. The hybrids showed better establishment ability (adaptability), precocity and higher yields than F3-Amazon [33, 34]. In Nigeria, the yield of Series II Hybrids could be higher than F3-Amazon yield by as much as 30% (165 kg/ha) [34].

*Second Breeding Phase (1961–1970):* The main focus during this period was on breeding of cocoa variety for specific ecological needs [35] with four primary objectives:

i.Select superior genotypes with high yield and desirable commercial qualities.

ii.Drought resistance or establishment ability

iii.*Phytophthora* pod rot resistance

iv.Cocoa swollen shoot virus (CSSV) resistance or tolerance

Two major outputs of this programme include selection of 12 "CRIN Establishment Ability" Elites which had a significant and prolonged experience in the south western part of Nigeria due to increased deforestation. A large number of germplasm was also introduced from Trinidad through a programme sponsored by the Cocoa Alliance, London [19].

*Third Breeding Phase (1971–1980):* The objective of this programme among others was to develop varieties that are resistant to or tolerant to pod rot disease caused by *Phytophthora megakarya* and *P. palmivora*. This programme was not fully implemented due to inadequate experts in the Breeding unit coupled with the challenge of self-incompatibility, a limitation to selfing of the Upper Amazon cacao trees [24]. Inadequate funding of breeding research was anther limiting factor.

*Fourth Breeding Phase (1998–2008):* This was a 10-year CFC/ICCO/IPGRI project with primary objective to develop new cacao varieties that will be high yielding, early bearing, resistant to Phytophthora pod rot, resistant to mirids and also have good and acceptable physical and chocolate flavor quality. The on-station and on-farm evaluations in this programme by CRIN resulted in the selection and consequent registration and release of eight new varieties (CRIN Tc-1, CRIN Tc-2, CRIN Tc-3, CRIN Tc-4, CRIN Tc-5, CRIN Tc-6, CRIN Tc-7 and CRIN Tc-8) in 2010. These hybrid varieties have diverse genetic base, they are early bearing, high yielding, with very low input, resistant to major pests and diseases of cacao, highly adaptable to cacao ecologies of Nigeria. The low input for high productivity attributes of these varieties is of great benefit for sustainable production of cocoa in Nigeria.

## **5. Current production challenges**

Although remarkable breeding efforts have been invested through germplasm acquisition and development of improved varieties with high yielding potentials, cocoa yield at the farm level in Nigeria is still low when compare with the yield

**111**

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria*

potentials of improved varieties and there is continuous decline in the national output over the past three decades. Sustainable production of cocoa is achievable when prior attention is given to all factors involved in the production line. The following key constraints have been reported by [8] to be responsible for the low yield

The solutions to most of these problems require intensive and focused breeding. Therefore funding cacao breeding research aimed at addressing these challenges should be given prompt attention. The role of the government in proffering solution to some of these problems cannot be over emphasized. Government would need to subsidize farm inputs needed for cocoa production. In addition to this there is the need for public private partnership efforts and intervention of stakeholders in cocoa

Recent advancement in the fields of genetics, breeding and biotechnology has been used to the benefit of cacao improvement worldwide. The first molecular markers used for cacao genetic diversity study were isozymes [36], but these have limited numbers of loci and low polymorphisms. Significant progress has been made in the past two decades in cacao genomic mapping and germplasm characterization as reviewed by Guiltinan et al. [5]. More attention needs to be given to genomic sciences of cacao since these tools could be used in addressing many of the unanswered questions in the area of yield, pests and diseases, drought, architecture and flavor quality for sustainability of cocoa production. One of the major challenges of cacao

**6. Advances in science for cocoa production sustainability**

i.In adequate supply of improved planting material

ii.Poor access to improved planting materials

*DOI: http://dx.doi.org/10.5772/intechopen.82703*

and production of cocoa in the country.

iii.Old age of trees

vi.Stem borer

vii.Mirid

xii.Termites

xiii.Bryophyte

iv.Black pod disease

viii.High cost of labour

v.Poor price of cocoa beans

ix.High price of chemicals and inputs

x.Loss of soil fertility/poor soil

xi.Adulterated chemicals

industry to address these challenges.

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria DOI: http://dx.doi.org/10.5772/intechopen.82703*

potentials of improved varieties and there is continuous decline in the national output over the past three decades. Sustainable production of cocoa is achievable when prior attention is given to all factors involved in the production line. The following key constraints have been reported by [8] to be responsible for the low yield and production of cocoa in the country.


iii.Old age of trees


Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

than F3-Amazon yield by as much as 30% (165 kg/ha) [34].

iv.Cocoa swollen shoot virus (CSSV) resistance or tolerance

Inadequate funding of breeding research was anther limiting factor.

Two major outputs of this programme include selection of 12 "CRIN Establishment Ability" Elites which had a significant and prolonged experience in the south western part of Nigeria due to increased deforestation. A large number of germplasm was also introduced from Trinidad through a programme sponsored by

*Third Breeding Phase (1971–1980):* The objective of this programme among others was to develop varieties that are resistant to or tolerant to pod rot disease caused by *Phytophthora megakarya* and *P. palmivora*. This programme was not fully implemented due to inadequate experts in the Breeding unit coupled with the challenge of self-incompatibility, a limitation to selfing of the Upper Amazon cacao trees [24].

*Fourth Breeding Phase (1998–2008):* This was a 10-year CFC/ICCO/IPGRI project with primary objective to develop new cacao varieties that will be high yielding, early bearing, resistant to Phytophthora pod rot, resistant to mirids and also have good and acceptable physical and chocolate flavor quality. The on-station and on-farm evaluations in this programme by CRIN resulted in the selection and consequent registration and release of eight new varieties (CRIN Tc-1, CRIN Tc-2, CRIN Tc-3, CRIN Tc-4, CRIN Tc-5, CRIN Tc-6, CRIN Tc-7 and CRIN Tc-8) in 2010. These hybrid varieties have diverse genetic base, they are early bearing, high yielding, with very low input, resistant to major pests and diseases of cacao, highly adaptable to cacao ecologies of Nigeria. The low input for high productivity attributes of these varieties is of great benefit for sustainable production of cocoa

Although remarkable breeding efforts have been invested through germplasm acquisition and development of improved varieties with high yielding potentials, cocoa yield at the farm level in Nigeria is still low when compare with the yield

ii.Drought resistance or establishment ability

iii.*Phytophthora* pod rot resistance

the Cocoa Alliance, London [19].

**5. Current production challenges**

*Second Breeding Phase (1961–1970):* The main focus during this period was on breeding of cocoa variety for specific ecological needs [35] with four primary

i.Select superior genotypes with high yield and desirable commercial qualities.

of the eleven approved Amazon T (Trinidad) types having broad parental background and have been shown to be superior to West African Amelonado in establishment ability, vegetative vigor, precocity (earliness in bearing) and yield. F3-Amazon variety is also tolerant of CSSV and capsid attack [23, 32]. The Mixed Series II Hybrids (WACRI Series II) varieties were produced from crosses of Upper Amazon cacao with local selections. The hybrids showed better establishment ability (adaptability), precocity and higher yields than F3-Amazon [33, 34]. In Nigeria, the yield of Series II Hybrids could be higher

**110**

in Nigeria.

objectives:


The solutions to most of these problems require intensive and focused breeding. Therefore funding cacao breeding research aimed at addressing these challenges should be given prompt attention. The role of the government in proffering solution to some of these problems cannot be over emphasized. Government would need to subsidize farm inputs needed for cocoa production. In addition to this there is the need for public private partnership efforts and intervention of stakeholders in cocoa industry to address these challenges.

## **6. Advances in science for cocoa production sustainability**

Recent advancement in the fields of genetics, breeding and biotechnology has been used to the benefit of cacao improvement worldwide. The first molecular markers used for cacao genetic diversity study were isozymes [36], but these have limited numbers of loci and low polymorphisms. Significant progress has been made in the past two decades in cacao genomic mapping and germplasm characterization as reviewed by Guiltinan et al. [5]. More attention needs to be given to genomic sciences of cacao since these tools could be used in addressing many of the unanswered questions in the area of yield, pests and diseases, drought, architecture and flavor quality for sustainability of cocoa production. One of the major challenges of cacao

**Figure 4.** *Cacao establishment on well irrigated land without the use of plantain shade.*

breeding is its long gestation period which takes a minimum of 2–3 years (from seed to seed). Cacao breeding is yet to tap from the advantage of marker assisted selection in reducing the breeding cycle as it is applicable in many other crops.

Tissue culture technique is useful not only for cryo preservation of germplasm materials. Application of somatic embryogenesis through temporary immersion technology will enhance mass clonal production of improved seedlings for large scale distribution to farmers to solve the problem of insufficient planting materials. Recent advancement in cacao science have been reported [37] that makes it possible for cacao planted on well irrigated land to survive without the conventional use of plantain as shade crop (**Figure 4**). This technology has the potential to solve the challenge of deforestation associated with new cacao establishment. This will also help to extend Nigeria cacao growing ecology to savannah region for increased productivity.

## **7. Conclusions**

The sustainability and future of Nigeria cocoa production is hinged on the amount of diversity of genetic resources conserved and utilized in development of planting materials for farmer. Funding of cacao germplasm collections and research on its conservation, evaluation and use for breeding should be a top priority and collective efforts of public private partnership. There is the need for targeted exploitation of useful underutilized genetic resources available in the germplasm collections for varietal development in future breeding program. Conservation of cacao genetic materials in Nigeria needs re-organization and efficient re-hauling by establishment of correctly identified clones in new breeders' core collection germplasm plot using technologies of barcoding labeling and drip irrigation systems. Cocoa production in Nigeria will be revived to attain sustainability if the sector could tap from the great potentials of scientific innovations and technological advancement to the advantage of the industry.

## **Acknowledgements**

The authors acknowledge and thank the Director/Chief Executive and the management of CRIN for granting the permission to publish this work.

**113**

**Author details**

Calabar, Nigeria

Festus Olakunle Olasupo1

provided the original work is properly cited.

© 2019 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

\* and Peter O. Aikpokpodion2

1 Plant Breeding Section, Cocoa Research Institute of Nigeria, Ibadan, Nigeria

2 Department of Genetics and Biotechnology, University of Calabar,

\*Address all correspondence to: festusolasupo@gmail.com

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria*

The authors declared that they have no conflict of interest.

*DOI: http://dx.doi.org/10.5772/intechopen.82703*

**Conflict of interest**

*Cacao Genetic Resources Conservation and Utilization for Sustainable Production in Nigeria DOI: http://dx.doi.org/10.5772/intechopen.82703*

## **Conflict of interest**

Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

*Cacao establishment on well irrigated land without the use of plantain shade.*

in reducing the breeding cycle as it is applicable in many other crops.

breeding is its long gestation period which takes a minimum of 2–3 years (from seed to seed). Cacao breeding is yet to tap from the advantage of marker assisted selection

Tissue culture technique is useful not only for cryo preservation of germplasm materials. Application of somatic embryogenesis through temporary immersion technology will enhance mass clonal production of improved seedlings for large scale distribution to farmers to solve the problem of insufficient planting materials. Recent advancement in cacao science have been reported [37] that makes it possible for cacao planted on well irrigated land to survive without the conventional use of plantain as shade crop (**Figure 4**). This technology has the potential to solve the challenge of deforestation associated with new cacao establishment. This will also help to extend Nigeria cacao growing ecology to savannah region for increased productivity.

The sustainability and future of Nigeria cocoa production is hinged on the amount of diversity of genetic resources conserved and utilized in development of planting materials for farmer. Funding of cacao germplasm collections and research on its conservation, evaluation and use for breeding should be a top priority and collective efforts of public private partnership. There is the need for targeted exploitation of useful underutilized genetic resources available in the germplasm collections for varietal development in future breeding program. Conservation of cacao genetic materials in Nigeria needs re-organization and efficient re-hauling by establishment of correctly identified clones in new breeders' core collection germplasm plot using technologies of barcoding labeling and drip irrigation systems. Cocoa production in Nigeria will be revived to attain sustainability if the sector could tap from the great potentials of scientific innovations and technological

The authors acknowledge and thank the Director/Chief Executive and the

management of CRIN for granting the permission to publish this work.

**112**

**Figure 4.**

**7. Conclusions**

**Acknowledgements**

advancement to the advantage of the industry.

The authors declared that they have no conflict of interest.

## **Author details**

Festus Olakunle Olasupo1 \* and Peter O. Aikpokpodion2

1 Plant Breeding Section, Cocoa Research Institute of Nigeria, Ibadan, Nigeria

2 Department of Genetics and Biotechnology, University of Calabar, Calabar, Nigeria

\*Address all correspondence to: festusolasupo@gmail.com

© 2019 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

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[3] Bartley BGD. The Genetic Diversity of Cacao and its Utilization. Wallingford: CABI; 2005. p. 341

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[10] GRA W, Lass RA. Cocoa. New York: Longman Scientific & Technical; 1985

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[13] Webster JB. The bible and the plough. Journal of Historical Society of Nigeria II. 1964;**4**:418-434

[14] Ayorinde JA. Historical notes on the introduction and development of the cocoa industry in Nigeria. Nigerian Agricultural Journal. 1966;**3**(1):18-23

[15] Berry SS. Christianity and the rise of cocoa growing in Ibadan and Ondo. Journal of the historical society of Nigeria IV. 1968;**3**:339-451

[16] Aikpokpodion PO, Motamayor JC, Adetimirin VO, Adu-Ampomah Y, Ingelbrecht I, Eskes AB, et al. Genetic diversity assessment of sub-samples of cacao, *Theobroma cacao* L. collections in West Africa using simple sequence repeats marker. Tree Genetics & Genomes. 2009;**5**:699-711

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*DOI: http://dx.doi.org/10.5772/intechopen.82703*

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MJ, Eskes A, Motamayor JC, et al. Population structure and molecular characterization of Nigerian field genebank collections of cacao, *Theobroma cacao* L. Silvae Genetica. 2010;**59**:273-285

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[21] Pound FJ. Cacao and witches' broom disease (*Marasmus perniciosus*). A report on a recent visit to the Amazon territory of Peru, September 1942–February 1943. Department of Agriculture, Trinidad and Tobago; 1943. 14 p

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[23] Toxopeus H. F3 Amazon cacao in Nigeria. Report of the Cocoa Research Institute of Nigeria. 1964. pp. 13-22

[24] Atanda OA. The Third Nigerian Cocoa Breeding Programme: A progress report 1977. In: 5th International Cocoa Research Conference Ibadan. 1977. pp. 85-90

[25] Olatoye ST, Esan EB. Recent innovation in cacao genetic resources conservation in Nigeria. In: Proceedings of the International Workshop on Conservation, Characterization and Utilization of Cacao Genetic Resources in the 21st Century. Cocoa Research Unit, Port of Spain, Trinidad 13-17 September, 1992. pp. 281-291

[26] Eskes AB, Efron Y, editors. Global Approaches to Cocoa Germplasm Utilization and Conservation. Final Report of the CFC/ICCO/IPGRI Project on 'Cocoa Germplasm utilization and Conservation: A Global Approach' (1998-2004). 2006. Amsterdam, The Netherlands: CFC; (ICCO, London, UK; and IPGRI, Rome, Italy)

[27] Aikpokpodion P. Genetic Diversity in Nigerian Cacao, (*Theobroma cacao* L.) Collections as Revealed by Phenotypic and Simple Sequence Repeats Marker. Ibadan, Nigeria: The University of Ibadan, Nigeria; 2007 131 p

[28] Olasupo FO, Adewale BA, Aikpokpodion PO, Muyiwa AA, Bhattacharjee R, Gutierrez OA, et al. Genetic identity and diversity of Nigerian cacao genebank collections verified by single nucleotide polymorphisms (SNPs): A guide to field genebank management and utilization. Tree Genetics & Genomes. 2018;**14**(32):1-16

[29] Cervantes-Martinez C, Brown JS, Schnell RJ, Phillips-Mora W, Takrama JF, Motamayor JC. Combining ability for disease resistance, yield and horticultural traits of cacao (*Thoebroma cacao* L.) clones. Journal of the American Society for Horticultural Science. 2006;**131**:231-241

[30] Takrama J, Kun J, Meinharddt L, Mischke S, Opoku S, Padi FK, et al. Verification of genetic identity of introduced cacao germplasm in Ghana using single nucleotide polymorphism (SNP) markers. African Journal of Biotechnology. 2014;**13**(2):127-136

**114**

Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

(*Theobroma cacao* L.) varieties and farm productivity in Nigeria. Journal of Agricultural Science. 2011;**3**(4):68-76

[9] Toxopeus H. History of the Nigerian cocoa crop in the light of the socioeconomic revolution of the 19th century in West Africa. Landbouwkundig Tijschrift. 1973;**83**(12):485-490

[10] GRA W, Lass RA. Cocoa. New York: Longman Scientific & Technical; 1985

[11] Lanaud C, Kébé I, Risterucci A, Clément D, N'Goran J, et al. Mapping quantitative trait loci (QTL) for resistance to *Phytophthora palmivora* in *T. Cacao*. Proceedings International

[12] Opeke LK. Development of cacao industry in Nigeria. In: Proceedings of the 6th Conference of the Agricultural Society of Nigeria. 1969. pp. 1-5

[13] Webster JB. The bible and the plough. Journal of Historical Society of

[14] Ayorinde JA. Historical notes on the introduction and development of the cocoa industry in Nigeria. Nigerian Agricultural Journal. 1966;**3**(1):18-23

[15] Berry SS. Christianity and the rise of cocoa growing in Ibadan and Ondo. Journal of the historical society of

[16] Aikpokpodion PO, Motamayor JC, Adetimirin VO, Adu-Ampomah Y, Ingelbrecht I, Eskes AB, et al. Genetic diversity assessment of sub-samples of cacao, *Theobroma cacao* L. collections in West Africa using simple sequence repeats marker. Tree Genetics & Genomes. 2009;**5**:699-711

[17] Aikpokpodion PO, Kolesnikova-Allen M, Adetimirin VO, Guiltinan

Nigeria II. 1964;**4**:418-434

Nigeria IV. 1968;**3**:339-451

Cocoa Research Conference.

2000;**12**:99-105

**References**

[1] Alverson WS, Whitlock BA, Nyfeller R, Bayer C, Baum DA. Phylogeny of the core *Malvales*: Evidence from *ndhF* sequence data. American Journal of

Botany. 1999;**86**:1474-1486

[2] Cheesman EE. Notes on nomenclature, classification and possible relationships of cacao populations. Tropical Agriculture.

[3] Bartley BGD. The Genetic

Diversity of Cacao and its Utilization. Wallingford: CABI; 2005. p. 341

[4] ICCO Production Values. 2018. Available from: http://www.icco.org/ faq/57.cocoa-production/123 [Accessed:

[5] Guiltinan MJ, Verica J, Zhang D, Figuerira A. Genomics of *Theobroma cacao*, "the food of the gods". In: Moore PH, Ming R, editors. Genomics of Tropical Crop Plants. Switzerland AG:

[6] Aikpokpodion PO. Variation in agro-morphological characteristics of cacao. *Theobroma cacao* L., in farmers' fields in Nigeria. New Zealand Journal of Crop and Horticultural Science. 2010;**38**(2):157-170. DOI: 10.1080/0028825X.2010.488786

[7] Aikpokpodion PO. Defining Genetic Diversity in the Chokolate Tree, *Theobroma cacao* L. Grown in West and Central Africa. In: Genetic Diversity in Plants. London, UK: InTech; 2012. ISBN: 978-51-0185-7. Available from: http://www.intechchopen.com/ books/genetic-diversity-in-plants/ defining-genetic-diversity-in-chocolatetree-thoebroma-cacao-l-grown-in-westand-central-africa [Accessed: Jul 28, 2014]

[8] Aikpokpodion PO, Adeogun SO. A diagnostic study of constraints to achieving yield potentials of cocoa

Springer; 2008. pp. 145-170

1944;**21**:144-159

Sep 15, 2018]

Chapter 7

Abstract

1. Introduction

117

Subbian Elain Apshara

Cocoa Genetic Resources and

Cropping Systems of India

Their Utilization in Palm-Based

Cocoa (Theobroma cacao L.) became an integral part of palm-based cropping systems of India. It is being grown profitably as a mixed crop under arecanut (Areca catechu L.), coconut (Cocos nucifera L.), and oil palm (Elaeis guineensis Jacq.) gardens of the southern states Karnataka, Kerala, Tamil Nadu, and Andhra Pradesh. It is well adapted to the humid tropics with high rainfall and short dry spells as well as irrigated gardens of tropical belts, utilizing the shade provided by the palms. Research efforts of more than four decades at the ICAR-CPCRI (Indian Council of Agricultural Research-Central Plantation Crops Research Institute) and KAU (Kerala Agriculture University) have allowed efficient utilization of conserved cocoa genetic resources by farmers to provide additional income through

multispecies cropping systems in the spices and plantation sector. National Horticulture Mission of Govt. of India identified cocoa as a potential crop for area expansion and development to meet both internal and export demands. Improved varieties were developed with high pod yield, bean quality, suitable to different agro-climatic zones and to tackle major biotic and abiotic stress. This chapter outlines the contributions of cocoa breeding efforts at the research institutes and State Agricultural Universities; developmental programs of Ministry of Agriculture and procurement and processing facilities to the growth of the cocoa sector in India.

Keywords: cacao, cocoa, dry beans, genetic resources, germplasm, breeding

Cocoa was introduced into India way back in 1798 at Courtallam in Tirunelveli District of the Old Madras State (Tamil Nadu) by East India Company [1]. Cocoa was distributed then in the agro-climatic region covering Western Ghats Hills and Plains of Malabar (Kerala) and Mysore (Karnataka) states, having more rainy days and short dry periods [2]. Commercial cultivation of cocoa as a plantation crop under palms started in early 1970s, and the current area under cocoa is 82,940 ha with a production of 18,920 tonnes of cocoa beans [3]. From the traditional hilly regions, cocoa production has shifted and expanded to coconut gardens of nontraditional areas of Tamil Nadu and Andhra Pradesh states utilizing the 50% shade available in the gardens and irrigation. Safe conservation of genetic resources of this introduced crop and its utilization in breeding program is a top priority considering its perennial nature, adaptability, and compatibility with palms. Cocoa tree has a

[31] Padi FK, Ofori A, Takrama J, Djan E, Opoku SY, Dadzie AM, et al. The impact of SNP fingerprinting and parentage analysis on the effectiveness of variety recommendations in cacao. Tree Genetics & Genomes. 2015;**11**:1-14

[32] Longworth JF. The effect of cocoa swollen shoot on mature cocoa. Tropical Agriculture. 1963;**40**:275-283

[33] Atanda OA. Assessment of WACRI series I and II varieties of cocoa in Nigeria 2. Pod value, butter fat content and flavor assessment. Ghana Journal of Agricultural Science. 1975;**86**:1474-1486

[34] Atanda OA, Jacobs VJ. Assessment of WACRI series I and II varieties of cocoa in Nigeria 1. Establishment and yield. Ghana Journal of Agricultural Science. 1974;**7**:215-220

[35] Atanda OA. Breeding cacao in Nigeria. Cacao Growers Bulletin. 1973;**20**:17-23

[36] Lanaud C. Genetic studies of *Theobroma cacao* L. with the help of enzymatic markers. I. Genetic control and linkage of nine enzymatic markers. Café Cacao Thé. 1986;**30**:259-270

[37] World Cocoa Foundation. The effect of science in cocoa production. Available from: http:// www.worldcocoafoundation. org/wp-content/uploads/ file\_mf/1382644618DAY2Artand scienceofcocoabreeding [Accessed: Oct 15, 2018]

## Chapter 7

Theobroma cacao *- Deploying Science for Sustainability of Global Cocoa Economy*

[31] Padi FK, Ofori A, Takrama J, Djan E, Opoku SY, Dadzie AM, et al. The impact of SNP fingerprinting and parentage analysis on the effectiveness of variety recommendations in cacao. Tree Genetics & Genomes. 2015;**11**:1-14

[32] Longworth JF. The effect of cocoa swollen shoot on mature cocoa. Tropical

[33] Atanda OA. Assessment of WACRI series I and II varieties of cocoa in Nigeria 2. Pod value, butter fat content and flavor assessment. Ghana Journal of Agricultural Science. 1975;**86**:1474-1486

[34] Atanda OA, Jacobs VJ. Assessment of WACRI series I and II varieties of cocoa in Nigeria 1. Establishment and yield. Ghana Journal of Agricultural

[35] Atanda OA. Breeding cacao in Nigeria. Cacao Growers Bulletin.

[36] Lanaud C. Genetic studies of *Theobroma cacao* L. with the help of enzymatic markers. I. Genetic control and linkage of nine enzymatic markers. Café Cacao Thé. 1986;**30**:259-270

[37] World Cocoa Foundation. The effect of science in cocoa production. Available from: http:// www.worldcocoafoundation. org/wp-content/uploads/

file\_mf/1382644618DAY2Artand scienceofcocoabreeding [Accessed: Oct

Agriculture. 1963;**40**:275-283

Science. 1974;**7**:215-220

1973;**20**:17-23

15, 2018]

**116**

## Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

Subbian Elain Apshara

## Abstract

Cocoa (Theobroma cacao L.) became an integral part of palm-based cropping systems of India. It is being grown profitably as a mixed crop under arecanut (Areca catechu L.), coconut (Cocos nucifera L.), and oil palm (Elaeis guineensis Jacq.) gardens of the southern states Karnataka, Kerala, Tamil Nadu, and Andhra Pradesh. It is well adapted to the humid tropics with high rainfall and short dry spells as well as irrigated gardens of tropical belts, utilizing the shade provided by the palms. Research efforts of more than four decades at the ICAR-CPCRI (Indian Council of Agricultural Research-Central Plantation Crops Research Institute) and KAU (Kerala Agriculture University) have allowed efficient utilization of conserved cocoa genetic resources by farmers to provide additional income through multispecies cropping systems in the spices and plantation sector. National Horticulture Mission of Govt. of India identified cocoa as a potential crop for area expansion and development to meet both internal and export demands. Improved varieties were developed with high pod yield, bean quality, suitable to different agro-climatic zones and to tackle major biotic and abiotic stress. This chapter outlines the contributions of cocoa breeding efforts at the research institutes and State Agricultural Universities; developmental programs of Ministry of Agriculture and procurement and processing facilities to the growth of the cocoa sector in India.

Keywords: cacao, cocoa, dry beans, genetic resources, germplasm, breeding

## 1. Introduction

Cocoa was introduced into India way back in 1798 at Courtallam in Tirunelveli District of the Old Madras State (Tamil Nadu) by East India Company [1]. Cocoa was distributed then in the agro-climatic region covering Western Ghats Hills and Plains of Malabar (Kerala) and Mysore (Karnataka) states, having more rainy days and short dry periods [2]. Commercial cultivation of cocoa as a plantation crop under palms started in early 1970s, and the current area under cocoa is 82,940 ha with a production of 18,920 tonnes of cocoa beans [3]. From the traditional hilly regions, cocoa production has shifted and expanded to coconut gardens of nontraditional areas of Tamil Nadu and Andhra Pradesh states utilizing the 50% shade available in the gardens and irrigation. Safe conservation of genetic resources of this introduced crop and its utilization in breeding program is a top priority considering its perennial nature, adaptability, and compatibility with palms. Cocoa tree has a

typical growth habit and a distinct morphology highly responsive to climate change and growing environments, which necessitates long-term conservation of genetic resources and dynamic breeding programs [4] as systematically adopted in India.

ICQC, R and collected exotic clones with desirable traits for specific research purpose, for which ICAR-NBPGR (National Bureau of Plant Genetic Resources) is the nodal agency in India. Around 500 collections are being maintained in the National Active Germplasm Site (NAGS) for cocoa at CPCRI, Regional Station, Vittal, Karnataka and also at KAU, Kerala. The germplasm collections include clones from The Amazon, Brazil, Ecuador, Ghana, Kew, Jamaica, Mexico, Nigeria, Peru, and local collections from Kallar in Tamil Nadu, Wayanad, Idukki in Kerala, and Shiradi

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

adaptability, precocity, compatibility, stability of yield, productivity, and quality of

Diversity among the genetic resources is important for improvement program, and Bartley [12] explained the existence of diversity based on the degree of human involvement in establishment of cocoa groups. The three basic types of cocoa, Criollo, Forastero, and Trinitario, which have specific pod and bean characteristics [13] are also among the collections. Expression of diversity is estimated from different indicators of variability, especially, morphological traits that are important for cataloging and characterization of germplasm. Bioversity International has standardized the descriptor status for cocoa, which comprises of 60 characteristics. Turnbull and Eskes [14] developed visual aid to identify widely distributed cocoa accessions with a minimal descriptor of 20 characters. Morphological variability with regard to tree architecture, leaves, flowers, fruit shapes, apex form, pod rugosity, prominence of ridges and furrows, husk thickness, pod size, color, bean size, shape, and color are characterized, and passport data documentation has been undertaken in 30-year old cocoa collections [15]. National identity numbers (Indigenous/Exotic Collection, IC/EC No.) were obtained from NBPGR, New Delhi for further utilization in the breeding program. Assessment of distinctness, uniformity

The presence of genetic variability offers wide scope in selection of potential types. Yield is the main selection criterion. An easy approach in yield improvement is to select superior plants and subsequently developing them into clones. The major selection criteria followed in cocoa are, trees yielding 100 pods/tree/year, pods weighing 350–400 g or more with 35–40 beans having a fermented single dry bean weight of 1 g. Dry bean production is in general considered as a combination of three yield components: bean weight per pod, number of pods per tree, and number of trees per hectare. It is expected to be 1 kg and above per tree in a year, and productivity is usually assessed over 6 years in varietal trials after stabilization. At CPCRI, seven high yielding clones VTLC-1, VTLC-5, VTLC-7, VTLC-8, VTLC-9, VTLC-11, and VTLC-30 were selected and utilized as parents in the breeding programs as well as in establishment of seed gardens [16]. KAU identified seven clones CCRP 1 to 7 and released these selections for cultivation. Though individual tree selections are being made from seedling progenies, they have to be further evaluated in clonal trials for confirmation. Heritability for yield is low or average when based on single tree harvests but higher when based on yields from plots containing several trees, and so clonal varieties are gaining importance [17]. To assess the phenotypic value of genotype even in hybrid selection programs, clonal trials are advised [18]. From the clonal trials, three varieties VTLCC-1, VTLCS-1, and VTLCS-2 have been released for commercial cultivation in different agro-climatic zones in the countryn (Table 1). Genetic analysis on 17 plant, pod, and bean characters in 44 exotic cocoa clones resulted in the selection of superior genotypes for higher performance with traits of high heritability and genetic advance. Based

Ghats of Karnataka. All these are being conserved and evaluated for their

produce in the introduced environment.

DOI: http://dx.doi.org/10.5772/intechopen.82077

and stability (DUS) of traits is currently underway.

3.2 Breeding through selection

119

## 2. History of cocoa improvement in India

Cocoa breeding is one of the oldest improvement programs in the world. In India, the oldest plantations with Criollo type cocoa were established and evaluated for their performance in Kallar and Burliar Fruit Stations in the Nilgiris between 1930 and 1935. Those found with high yield potentials were distributed wherever the agro-climatic conditions suited the crop. In 1962, Indian Council of Agricultural Research (ICAR) decreed to grow Criollo type in South India and Forastero type in North East India. In 1964, few Malaysian clones of Forastero and Trinitario types were imported, and research on arecanut + cocoa and coconut + cocoa mixed cropping systems were conducted at Central Plantation Crops Research Institute (CPCRI), Regional Station, Vittal, Karnataka and other centers, Peechi and Palode in Kerala and Kahikuchi in Assam and proved as profitable cropping models. In 1965, a research-cum-demonstration unit of Cadbury India Pvt. Ltd. was established in Chundale in Kerala [5]. In 1969, systematic research was started in CPCRI with additional introductions of germplasm, followed by Kerala Agricultural University (KAU) in 1979 and then continued at KAU in 1987 with Cadbury India Pvt. Ltd. funding. In 2008, Tamil Nadu Agricultural University (TNAU) initiated cocoa research with funding from Mondelez International.

## 3. Breeding strategies in India

Systematic and long-term cocoa improvement programs are being taken up with the following strategies: (1) germplasm collection, conservation, cataloging, characterization, and evaluation; (2) breeding through selection and hybridization; (3) standardization of vegetative multiplication, establishment of seed gardens/clonal orchards; (4) comparative yield trials (CYT); (5) multi-locational trials (MLT); (6) screening and resistance breeding for biotic and abiotic stress; (7) biotechnology and bioinformatics approaches; and (8) demonstration gardens [6, 7].

#### 3.1 Introduction of germplasm

The basic step in any breeding program is the introduction of germplasm from both the primary and secondary centers of origin and distribution. Cocoa in its native zones of South and Central America and other major producing countries of Africa is suffering with many debilitating diseases caused by both viral and fungal pathogens and serious pests. To safe guard the germplasm exchange program, Intermediate Cocoa Quarantine Centre (ICQC, R) was established in Europe. The University of Reading took over the responsibility for cocoa quarantine in 1987 [8]. The center routinely collects important clones from international and national gene banks, wild collections and conducts virus indexing. It conducts strict quarantine for other major diseases and pests using the guidelines for safe movement of germplasm formulated by FAO/Bioversity International [9, 10]. Clones that have been cleared at the quarantine are supplied as bud sticks, with proper import permits and sanitary certificates to cocoa researchers over 20 countries. It also maintains the International Cocoa Germplasm Database to know about the gene banks, clones, traits, and even SSR/SNP profiles [11]. CPCRI also utilizes facilities offered by

### Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

ICQC, R and collected exotic clones with desirable traits for specific research purpose, for which ICAR-NBPGR (National Bureau of Plant Genetic Resources) is the nodal agency in India. Around 500 collections are being maintained in the National Active Germplasm Site (NAGS) for cocoa at CPCRI, Regional Station, Vittal, Karnataka and also at KAU, Kerala. The germplasm collections include clones from The Amazon, Brazil, Ecuador, Ghana, Kew, Jamaica, Mexico, Nigeria, Peru, and local collections from Kallar in Tamil Nadu, Wayanad, Idukki in Kerala, and Shiradi Ghats of Karnataka. All these are being conserved and evaluated for their adaptability, precocity, compatibility, stability of yield, productivity, and quality of produce in the introduced environment.

Diversity among the genetic resources is important for improvement program, and Bartley [12] explained the existence of diversity based on the degree of human involvement in establishment of cocoa groups. The three basic types of cocoa, Criollo, Forastero, and Trinitario, which have specific pod and bean characteristics [13] are also among the collections. Expression of diversity is estimated from different indicators of variability, especially, morphological traits that are important for cataloging and characterization of germplasm. Bioversity International has standardized the descriptor status for cocoa, which comprises of 60 characteristics. Turnbull and Eskes [14] developed visual aid to identify widely distributed cocoa accessions with a minimal descriptor of 20 characters. Morphological variability with regard to tree architecture, leaves, flowers, fruit shapes, apex form, pod rugosity, prominence of ridges and furrows, husk thickness, pod size, color, bean size, shape, and color are characterized, and passport data documentation has been undertaken in 30-year old cocoa collections [15]. National identity numbers (Indigenous/Exotic Collection, IC/EC No.) were obtained from NBPGR, New Delhi for further utilization in the breeding program. Assessment of distinctness, uniformity and stability (DUS) of traits is currently underway.

## 3.2 Breeding through selection

The presence of genetic variability offers wide scope in selection of potential types. Yield is the main selection criterion. An easy approach in yield improvement is to select superior plants and subsequently developing them into clones. The major selection criteria followed in cocoa are, trees yielding 100 pods/tree/year, pods weighing 350–400 g or more with 35–40 beans having a fermented single dry bean weight of 1 g. Dry bean production is in general considered as a combination of three yield components: bean weight per pod, number of pods per tree, and number of trees per hectare. It is expected to be 1 kg and above per tree in a year, and productivity is usually assessed over 6 years in varietal trials after stabilization. At CPCRI, seven high yielding clones VTLC-1, VTLC-5, VTLC-7, VTLC-8, VTLC-9, VTLC-11, and VTLC-30 were selected and utilized as parents in the breeding programs as well as in establishment of seed gardens [16]. KAU identified seven clones CCRP 1 to 7 and released these selections for cultivation. Though individual tree selections are being made from seedling progenies, they have to be further evaluated in clonal trials for confirmation. Heritability for yield is low or average when based on single tree harvests but higher when based on yields from plots containing several trees, and so clonal varieties are gaining importance [17]. To assess the phenotypic value of genotype even in hybrid selection programs, clonal trials are advised [18]. From the clonal trials, three varieties VTLCC-1, VTLCS-1, and VTLCS-2 have been released for commercial cultivation in different agro-climatic zones in the countryn (Table 1). Genetic analysis on 17 plant, pod, and bean characters in 44 exotic cocoa clones resulted in the selection of superior genotypes for higher performance with traits of high heritability and genetic advance. Based

typical growth habit and a distinct morphology highly responsive to climate change and growing environments, which necessitates long-term conservation of genetic resources and dynamic breeding programs [4] as systematically adopted in India.

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

Cocoa breeding is one of the oldest improvement programs in the world. In India, the oldest plantations with Criollo type cocoa were established and evaluated for their performance in Kallar and Burliar Fruit Stations in the Nilgiris between 1930 and 1935. Those found with high yield potentials were distributed wherever the agro-climatic conditions suited the crop. In 1962, Indian Council of Agricultural Research (ICAR) decreed to grow Criollo type in South India and Forastero type in North East India. In 1964, few Malaysian clones of Forastero and Trinitario types were imported, and research on arecanut + cocoa and coconut + cocoa mixed cropping systems were conducted at Central Plantation Crops Research Institute (CPCRI), Regional Station, Vittal, Karnataka and other centers, Peechi and Palode in Kerala and Kahikuchi in Assam and proved as profitable cropping models. In 1965, a research-cum-demonstration unit of Cadbury India Pvt. Ltd. was

established in Chundale in Kerala [5]. In 1969, systematic research was started in CPCRI with additional introductions of germplasm, followed by Kerala Agricultural University (KAU) in 1979 and then continued at KAU in 1987 with Cadbury India Pvt. Ltd. funding. In 2008, Tamil Nadu Agricultural University (TNAU) initiated

Systematic and long-term cocoa improvement programs are being taken up with the following strategies: (1) germplasm collection, conservation, cataloging, characterization, and evaluation; (2) breeding through selection and hybridization; (3) standardization of vegetative multiplication, establishment of seed gardens/clonal orchards; (4) comparative yield trials (CYT); (5) multi-locational trials (MLT); (6) screening and resistance breeding for biotic and abiotic stress; (7) biotechnology and

The basic step in any breeding program is the introduction of germplasm from both the primary and secondary centers of origin and distribution. Cocoa in its native zones of South and Central America and other major producing countries of Africa is suffering with many debilitating diseases caused by both viral and fungal pathogens and serious pests. To safe guard the germplasm exchange program, Intermediate Cocoa Quarantine Centre (ICQC, R) was established in Europe. The University of Reading took over the responsibility for cocoa quarantine in 1987 [8]. The center routinely collects important clones from international and national gene banks, wild collections and conducts virus indexing. It conducts strict quarantine for other major diseases and pests using the guidelines for safe movement of germplasm formulated by FAO/Bioversity International [9, 10]. Clones that have been cleared at the quarantine are supplied as bud sticks, with proper import permits and sanitary certificates to cocoa researchers over 20 countries. It also maintains the International Cocoa Germplasm Database to know about the gene banks, clones, traits, and even SSR/SNP profiles [11]. CPCRI also utilizes facilities offered by

cocoa research with funding from Mondelez International.

bioinformatics approaches; and (8) demonstration gardens [6, 7].

3. Breeding strategies in India

3.1 Introduction of germplasm

118

2. History of cocoa improvement in India

## Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy


Cocoa butter with free fatty acid (FFA) content of 1% or less together with acceptable flavor is the best indication of good quality beans. The type of fatty acids in 18 hybrids and 10 elite clones has been assessed, and from the profile, it is now known that 11 fatty acids are involved in the quality of cocoa beans. The fatty acids palmitic, stearic, oleic, linoleic, and arachidic acids present in all the genotypes invariably. The percentage of stearic acid is the highest in a range of 30.5% in VTLCP-7 to 44.2% in VTLCP-1. The other fatty acids differed among the genotypes

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

Hybrid vigor between parents showing good combining ability is readily exploited in cocoa improvement programs along with inter-population heterosis. Most commonly adopted method is developing hybrids between two distant

For production of true hybrids with specific objectives and to confirm the compatibility reaction, hand pollination is routinely practiced. In artificial pollination, a flower bud, which will open the following day, recognized by its whitish color and swollen appearance, is selected. The bud is covered with a hood of plastic tube of 5 1.5–2.0 cm size, which is sealed to the bark using materials like plasticine/glaze putty. The tube is covered with muslin cloth at the top, kept in place with a rubber band. Opened flowers are collected from the desired male parent, and stamens are carefully taken out by pushing the corresponding petal. One entire anther with a part of the filament is deposited on the stigma. The pollinated flowers are labeled using tin foil pieces fixed in the cushion using ball pins. The hoods are removed 24 hours after pollination, and in 3–5 days, fertilization is confirmed by

Different cross combinations have been tried with specific objectives for development of hybrids. At CPCRI, five progeny trials have been evaluated with 76 cross combinations during 1983–1994 at Vittal, Kidu, and Kasaragod centers with objectives of more number of pods, high dry bean yield, big bean size, and drought tolerance. Of these, 20 hybrids were identified as best hybrids and further evaluated as clones. Among them, 17 progenies exhibited high vigor and cropping efficiency even at early years of development [27, 28]. From the progeny trials, four hybrids VTLCH-1, VTLCH-2, VTLCH-3, and VTLCH-4 have been released as improved varieties for cultivation in the country; in 2006, the Golden Jubilee year of CPCRI, RS, Vittal and VTLCH-5 is released as Nethra Centura for the centenary year of

Hybridization program at KAU was initiated during 1983 and continued up to

1993. During this period, a total of 159 parents were included to produce 239 crosses, 187 pods, and 21,819 hybrid seedlings. Nursery evaluation of these hybrids is done with HD<sup>2</sup> (H—height and D—diameter) value. A total of 3126 superior seedlings were field planted in series I, II, III, and IV and progeny trials I, II, III, and IV, and 163 superior hybrids were selected, utilized in various breeding program for release of hybrids, and used as better combiners in clonal gardens. Three hybrids CCRP 8, CCRP 9, and CCRP 10 with high yield and good level of tolerance to

in the percentage of expressio.

genotypes.

3.3 Breeding through hybridization

DOI: http://dx.doi.org/10.5772/intechopen.82077

3.3.1 Method of hand pollination and fruit set

the visual swelling of the ovary [26].

CPCRI, Kasaragod (Tables 2 and 3).

3.3.2 Progeny trials

121

VTLCC—Vittal Cocoa Clone.

VTLCS—Vittal Cocoa Selection.

#### Table 1.

CPCRI cocoa varieties developed through selection.

on the pod yield, VTLC-25, VTLC-15, VTLC-18, VTLC-36, VTLC-13, VTLC-37, and VTLC-17 have been identified as heavy bearers with an optimal canopy. These clones recorded single dry bean weight of more than 1 g, 10–15% shell, high nib recovery 85–90%, and more than 50% butter fat content making them suitable for industries as well [19, 20].

In the palm-based inter cropping systems, the pod yield in general is expressed with respect to the canopy area which is mainly maintained as cone/ umbrella shaped single tier architecture. In the evaluation trials of Trinidad cocoa and Wayanad collections, 5 clones each are selected for high pod and dry bean yield ranging from 2.2 to 3.3 kg/tree/year [21, 22] with an optimal canopy of 15–20 m<sup>2</sup> . Trait specific improvements are being taken up in the current breeding programs. A bean index of 100 beans/100 g, that is, dry bean weight of 1 g is considered as a standard for grade I beans [23], and so selections are aimed at a larger bean size of 1 g and above, which will have high butter content suitable for chocolate industry. The bean size is influenced by genotype, environment, and also the processing methods. Box, basket, and tray methods are being examined by research institutes as well as by farmers. Variation in bean indices has been observed across collections and the single dry bean weight ranged from 0.7 to 2.5 g. To improve the qualitative parameters, Criollo cocoa is used in hybridization programs [24], and white bean genotypes are being evaluated for quality parameters. Cocoa is considered as functional food, and so dark chocolates are consumed for their health benefits. Catechin, epicatechin, and procyanidines are the main polyphenols present in cocoa contributing to bitterness, astringency, and the organoleptic quality of cocoa. Cocoa beans of different clones evaluated for polyphenols and antioxidant activity exhibited distinct differences. Polyphenols ranged from 82 to 136 mg/g, procyanidin 49 to 64 mg/g, fat content of 24–55%, and antioxidant activity of 77–98% among cocoa clones studied. In general, cocoa beans with high polyphenol and procyanidin contents exhibited high antioxidant activities which are utilized for qualitative improvement [25].

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

Cocoa butter with free fatty acid (FFA) content of 1% or less together with acceptable flavor is the best indication of good quality beans. The type of fatty acids in 18 hybrids and 10 elite clones has been assessed, and from the profile, it is now known that 11 fatty acids are involved in the quality of cocoa beans. The fatty acids palmitic, stearic, oleic, linoleic, and arachidic acids present in all the genotypes invariably. The percentage of stearic acid is the highest in a range of 30.5% in VTLCP-7 to 44.2% in VTLCP-1. The other fatty acids differed among the genotypes in the percentage of expressio.

### 3.3 Breeding through hybridization

Hybrid vigor between parents showing good combining ability is readily exploited in cocoa improvement programs along with inter-population heterosis. Most commonly adopted method is developing hybrids between two distant genotypes.

#### 3.3.1 Method of hand pollination and fruit set

For production of true hybrids with specific objectives and to confirm the compatibility reaction, hand pollination is routinely practiced. In artificial pollination, a flower bud, which will open the following day, recognized by its whitish color and swollen appearance, is selected. The bud is covered with a hood of plastic tube of 5 1.5–2.0 cm size, which is sealed to the bark using materials like plasticine/glaze putty. The tube is covered with muslin cloth at the top, kept in place with a rubber band. Opened flowers are collected from the desired male parent, and stamens are carefully taken out by pushing the corresponding petal. One entire anther with a part of the filament is deposited on the stigma. The pollinated flowers are labeled using tin foil pieces fixed in the cushion using ball pins. The hoods are removed 24 hours after pollination, and in 3–5 days, fertilization is confirmed by the visual swelling of the ovary [26].

#### 3.3.2 Progeny trials

on the pod yield, VTLC-25, VTLC-15, VTLC-18, VTLC-36, VTLC-13, VTLC-37, and VTLC-17 have been identified as heavy bearers with an optimal canopy. These clones recorded single dry bean weight of more than 1 g, 10–15% shell, high nib recovery 85–90%, and more than 50% butter fat content making them suitable for

Characters VTLCC 1 VTLCS 1 VTLCS 2

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

No. of pods/tree/year 55 55 55 No. of beans/pod 35 42 41 Single dry bean weight (g) 0.9–1.05 1.3 1.21 Dry bean yield kg/tree/year 1.3 2.5 2.7 Dry bean yield kg/ha 890 1700 1840 Shelling % 12 11 15 Nib recovery % 88 88 85 Fat content % 50 52 53

> suitable for North Eastern Zones

) 12 12 15

High yielder withstands both biotic and abiotic stress

High bean index, stable yielder withstands both biotic and abiotic stress

In the palm-based inter cropping systems, the pod yield in general is expressed with respect to the canopy area which is mainly maintained as cone/ umbrella shaped single tier architecture. In the evaluation trials of Trinidad cocoa and Wayanad collections, 5 clones each are selected for high pod and dry bean yield ranging from 2.2 to 3.3 kg/tree/year [21, 22] with an optimal canopy

. Trait specific improvements are being taken up in the current

breeding programs. A bean index of 100 beans/100 g, that is, dry bean weight of 1 g is considered as a standard for grade I beans [23], and so selections are aimed at a larger bean size of 1 g and above, which will have high butter content suitable for chocolate industry. The bean size is influenced by genotype, environment, and also the processing methods. Box, basket, and tray methods are being examined by research institutes as well as by farmers. Variation in bean indices has been observed across collections and the single dry bean weight ranged from 0.7 to 2.5 g. To improve the qualitative parameters, Criollo cocoa is used in hybridization programs [24], and white bean genotypes are being evaluated for quality parameters. Cocoa is considered as functional food, and so dark chocolates are consumed for their health benefits. Catechin, epicatechin, and procyanidines are the main polyphenols present in cocoa contributing to bitterness, astringency, and the organoleptic quality of cocoa. Cocoa beans of different clones evaluated for polyphenols and antioxidant activity exhibited distinct differences. Polyphenols ranged from 82 to 136 mg/g, procyanidin 49 to 64 mg/g, fat content of 24–55%, and antioxidant activity of 77–98% among cocoa clones studied. In general, cocoa beans with high polyphenol and procyanidin contents exhibited high antioxidant activities which are

industries as well [19, 20].

VTLCC—Vittal Cocoa Clone.

VTLCS—Vittal Cocoa Selection.

Features Self compatible line

CPCRI cocoa varieties developed through selection.

Canopy area (m<sup>2</sup>

utilized for qualitative improvement [25].

of 15–20 m<sup>2</sup>

120

Table 1.

Different cross combinations have been tried with specific objectives for development of hybrids. At CPCRI, five progeny trials have been evaluated with 76 cross combinations during 1983–1994 at Vittal, Kidu, and Kasaragod centers with objectives of more number of pods, high dry bean yield, big bean size, and drought tolerance. Of these, 20 hybrids were identified as best hybrids and further evaluated as clones. Among them, 17 progenies exhibited high vigor and cropping efficiency even at early years of development [27, 28]. From the progeny trials, four hybrids VTLCH-1, VTLCH-2, VTLCH-3, and VTLCH-4 have been released as improved varieties for cultivation in the country; in 2006, the Golden Jubilee year of CPCRI, RS, Vittal and VTLCH-5 is released as Nethra Centura for the centenary year of CPCRI, Kasaragod (Tables 2 and 3).

Hybridization program at KAU was initiated during 1983 and continued up to 1993. During this period, a total of 159 parents were included to produce 239 crosses, 187 pods, and 21,819 hybrid seedlings. Nursery evaluation of these hybrids is done with HD<sup>2</sup> (H—height and D—diameter) value. A total of 3126 superior seedlings were field planted in series I, II, III, and IV and progeny trials I, II, III, and IV, and 163 superior hybrids were selected, utilized in various breeding program for release of hybrids, and used as better combiners in clonal gardens. Three hybrids CCRP 8, CCRP 9, and CCRP 10 with high yield and good level of tolerance to

## Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy


#### Table 2.

Progeny trials of CPCRI.


#### Table 3.

CPCRI varieties developed through hybridization.

Vascular Streak Dieback (VSD) have been released as varieties for cultivation in the country (Table 4) [24].

established for production of hybrid seeds with known parentage and performance. These clonal orchards are established and maintained at CPCRI, Research Centre, Kidu, Nettana, Karnataka [30] and at College of Horticulture, KAU, Vellanikkara, Thrissur. In cocoa seed pods, seedlings and clones (grafts/budded plants) are being used as planting materials. CPCRI nurseries are accredited by National Horticulture Board (NHB), with four star rating for quality planting material supply and acting as model nursery on cocoa. Sixteen regional nurseries were established in different

states for the area expansion programs of Govt. of India.

Ref. [26].

Table 4.

123

Cocoa varieties of KAU.

Variety Breeding method Important traits

DOI: http://dx.doi.org/10.5772/intechopen.82077

CCRP-1 Single tree selection from local population Heavy bearer, self incompatible

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

CCRP-2 Selection Heavy bearer, self incompatible, VSD tolerant

CCRP-3 Selection Heavy bearer, self incompatible, VSD tolerant

CCRP-4 Selection Heavy bearer, self incompatible, VSD tolerant

CCRP-5 Selection Heavy bearer, self incompatible, VSD tolerant

CCRP-6 Selection Heavy bearer, self incompatible, VSD tolerant

CCRP-7 Selection Heavy bearer, self incompatible, VSD tolerant

CCRP-8 Hybridization High yielder and VSD tolerant

CCRP-9 Hybridization High yielder and VSD tolerant

CCRP-10 Hybridization High yielder and VSD tolerant

Vascular Streak Dieback (VSD) tolerant

No. of pods—56.2/tree/year No. of beans/pod—46.2 Single dry bean weight—0.8 g

No. of pods—53.9/tree/year No. of beans/pod—45.5 Single dry bean weight—1.0 g

No. of pods—68.5/tree/year No. of beans/pod—42.3 Single dry bean weight—1.0 g

No. of pods—66.2/tree/year No. of beans/pod—45.4 Single dry bean weight—1.1 g

No. of pods—37.9/tree/year No. of beans/pod—45.25 Single dry bean weight—0.8 g

No. of pods—50.1/tree/year No. of beans/pod—48 Single dry bean weight—1.9 g

No. of pods—78.1/tree/year No. of beans/pod—46.9 Single dry bean weight—0.9 g

No. of pods—90.4/tree/year No. of beans/pod—48.8 Single dry bean weight—0.9 g

No. of pods—106.7/tree/year No. of beans/pod—36.7 Single dry bean weight—0.8 g

No. of pods—79.6/tree/year No. of beans/pod—41.6 Single dry bean weight—1.1 g

#### 3.3.3 Seed gardens/clonal orchards

Based on the compatibility reactions, self-incompatible but cross-compatible high yielding parents are selected and multiplied as clones through soft wood grafting and established as clonal orchards or seed gardens. Bi-clonal and poly-clonal gardens were


Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

## Table 4.

Vascular Streak Dieback (VSD) have been released as varieties for cultivation in the

Suitable for water limited conditions

Trial Progenies tested Progenies selected Dry bean yield (kg/tree/year)

VTLCP-49 1.47 VTLCP-50 1.42 VTLCP-11 1.39

VTLCP-30 1.52

VTLCP-27 1.62

Progeny I (1983) 5 VTLCP-1 1.01 Progeny II (1984) 25 VTLCP-7 1.48

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

Progeny III (1987) 13 VTLCP-18 1.08 Progeny IV (1992) 15 VTLCP-29 1.25

Progeny V (1994) 18 VTLCP-26 1.33

Characters VTLCH 1 VTLCH 2 VTLCH 3 VTLCH 4 VTLCH 5

No. of beans/pod 40 40 41 40 43

Shelling % 13 11 15 15 11 Nib recovery % 87 89 87 87 88 Fat content % 54 54 51 51 52

> Heavy bearer, tolerant to black pod rot

16 15 18 18 16

50 50 41 40 66

1–1.11 1–1.5 1–1.05 1–1.07 1–1.11

1.4 1.5 1.7 1.6 2.5–3.0

959 1030 1150 1090 1500–1800

Suitable for water limited conditions

Suitable for high density planting both under arecanut and coconut

Based on the compatibility reactions, self-incompatible but cross-compatible high yielding parents are selected and multiplied as clones through soft wood grafting and established as clonal orchards or seed gardens. Bi-clonal and poly-clonal gardens were

country (Table 4) [24].

Table 3.

122

Features Early

VTLCH—Vittal Cocoa Hybrid [29].

stable heavy bearer

VTLCP—Vittal Cocoa Progeny [27].

Progeny trials of CPCRI.

Canopy area (m<sup>2</sup> )

of pods/tree/year

Single dry bean weight (g)

Dry bean yield kg/tree/year

Dry bean yield kg/ha

No.

Table 2.

3.3.3 Seed gardens/clonal orchards

CPCRI varieties developed through hybridization.

Cocoa varieties of KAU.

established for production of hybrid seeds with known parentage and performance. These clonal orchards are established and maintained at CPCRI, Research Centre, Kidu, Nettana, Karnataka [30] and at College of Horticulture, KAU, Vellanikkara, Thrissur. In cocoa seed pods, seedlings and clones (grafts/budded plants) are being used as planting materials. CPCRI nurseries are accredited by National Horticulture Board (NHB), with four star rating for quality planting material supply and acting as model nursery on cocoa. Sixteen regional nurseries were established in different states for the area expansion programs of Govt. of India.


demonstration plot scheme in farmers plots as well as in five AICRPP (All India Coordinated Research Project on Palms) centers funded by Directorate of Cashewnut and Cocoa Development (DCCD), Kochi and Directorate of Arecanut and Spices Development (DASD), Calicut, for identification of location specific varieties,

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

Chemical control can be effective against fungal diseases but will pollute the environment and make the cultivation expensive. Integrated disease and pest management by the use of resistant materials and cultural and biological methods is probably the best way to contain pathogens and pests in the long run for sustainable crop protection. Breeding for resistance, therefore, became the primary objective for cocoa breeders worldwide. Sources of resistance have been identified for major diseases, and since cocoa genome is sequenced, it is expected to provide models for plant pathogen interactions and also facilitate identification of resistance genes. Many cocoa pests such as mealy bugs, aphids, caterpillars, and borers are also reported in India [37] are currently being managed with chemical control measures. Screening of available germplasm for prevailing diseases, existing and emerging pests will therefore be very important in light of seasonal

India is free of most of the debilitating viral and fungal diseases known in cocoa. Since the current cocoa growing area comes under high rainfall zone and the main harvest season coincides with monsoon, incidence of black pod rot caused by Phytophthora palmivora is comparatively higher. Though the main harvest is safe guarded with systematic annual pruning, in the post monsoon period, the second season crop is still affected by pod rot. On field screening, clones have been categorized into having <10, 10–25, and >25% damage levels. In-vitro screening using isolates of P. palmivora, P. capsici, and P. citrophthora indicated that collections of Nigerian origin exhibit certain degree of tolerance [38]. Few Wayanad collections have also expressed field tolerance to pod rot when tested over three seasons. Further, 21 exotic clones collected exclusively for Phytophthora pod rot resistance was identified for utilization in cocoa hybridization program. The variety VTLCH-2, a combination of ICS 6 SCA 6 is found to be tolerant to black pod rot in India as well. KAU has taken up screening and hybridization program for combining desirable traits of CCRP released varieties and black pod

In India, vascular streak die back (VSD) caused by Oncobasidium theobromae was

first reported from Kottayam, Kerala, and it began to spread to adjoining cocoa growing areas of the state. As this disease cannot be controlled effectively by the use of fungicides, KAU breeding program concentrated mainly on production of VSD resistant varieties. Hybrid seedlings were screened in the nursery by subjecting them to high inoculum load by keeping them in the midst of infected seedlings. The tolerant and vigorous seedlings were selected and established in field for evaluation. CCRP varieties have been especially released for VSD resistance and also have been

utilized for establishment of clonal gardens for seedling supply [31].

common varieties, and to tackle the climate change effects.

3.6 Resistance breeding

DOI: http://dx.doi.org/10.5772/intechopen.82077

weather variations.

3.6.1 Black pod rot

resistance in cocoa.

125

3.6.2 Vascular streak die back

Table 5.

Inbred population field established (1989–2010) at KAU.

## 3.3.4 Inbreeding

Inbreeding forms a part of the breeding activities, not only to breed parents with some degree of homozygosity for the production of hybrids but also to breed materials homozygous for desirable traits like disease resistance. Existence of selfincompatibility makes inbreeding efforts in cocoa difficult. Since few selfcompatible trees are also identified in the populations, selfing is possible but it should be continued up to six to seven generations to attain homozygosity and thereafter to be utilized for crossing to exploit the hybrid vigor. KAU has taken up this task of selfing self-compatible plants and with over 20 years of continuous effort, maintains two genotypes of S4 generation, 5 of S3, 9 of S2, and 51 genotypes of S1 (Table 5) [24, 26, 31, 32].

## 3.4 Comparative yield trial (CYT)

The clones and progenies developed through selection and hybridization programs are multiplied as clones and evaluated under comparative yield trials in different situations. Under high density planting in arecanut garden, five hybrids VTLCP-6, VTLCP-20, VTLCP-15, VTLCP-1, and VTLCP-19 have been identified as best performers even in their initial years of growth [28]. Comparative study of parents and progenies as clones resulted in identification of VTLCP-6, VTLCP-2, and VTLCP-20 and parents VTLC-1 and VTLC-56 as potential high yielders [33]. In another trial, clones suitable for both arecanut and coconut canopies have been identified [34] and released as varieties. Under coconut in double hedge system of planting, hybrids VTLCP-22, VTLCP-18, and VTLCP-1 showed the best performance with an optimal canopy and high yield [35]. Evaluation of clonal and seedling progenies of selected genotypes has resulted in identification of four hybrids and two clones for multiplication both as clones and seedlings for utilization in the area expansion program [36].

## 3.5 Multi location trial (MLT) and demonstration plots

To assess the survival and stability of hybrids and clones in different agroclimatic conditions, multi-location trials are important. Elite clones of cocoa are under evaluation in both traditional and nontraditional states, namely Karnataka, Kerala, Tamil Nadu, Andhra Pradesh, Maharashtra, West Bengal, and Assam, for studying genotype location environment interactions. Further, 115 front line demonstration plots were established under participatory research cum

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

demonstration plot scheme in farmers plots as well as in five AICRPP (All India Coordinated Research Project on Palms) centers funded by Directorate of Cashewnut and Cocoa Development (DCCD), Kochi and Directorate of Arecanut and Spices Development (DASD), Calicut, for identification of location specific varieties, common varieties, and to tackle the climate change effects.

#### 3.6 Resistance breeding

Chemical control can be effective against fungal diseases but will pollute the environment and make the cultivation expensive. Integrated disease and pest management by the use of resistant materials and cultural and biological methods is probably the best way to contain pathogens and pests in the long run for sustainable crop protection. Breeding for resistance, therefore, became the primary objective for cocoa breeders worldwide. Sources of resistance have been identified for major diseases, and since cocoa genome is sequenced, it is expected to provide models for plant pathogen interactions and also facilitate identification of resistance genes. Many cocoa pests such as mealy bugs, aphids, caterpillars, and borers are also reported in India [37] are currently being managed with chemical control measures. Screening of available germplasm for prevailing diseases, existing and emerging pests will therefore be very important in light of seasonal weather variations.

#### 3.6.1 Black pod rot

3.3.4 Inbreeding

Table 5.

of S1 (Table 5) [24, 26, 31, 32].

area expansion program [36].

124

3.5 Multi location trial (MLT) and demonstration plots

3.4 Comparative yield trial (CYT)

Inbred population field established (1989–2010) at KAU.

Inbreeding forms a part of the breeding activities, not only to breed parents with

Generation Genotypes Plants selfed Flowers selfed Pods obtained S0 102 102 25,052 147 S1 51 178 6263 163 S2 9 41 693 24 S3 5 55 1720 48 S4 2 17 428 9 S5 1 1 132 0 Total 75 394 34,288 391

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

The clones and progenies developed through selection and hybridization programs are multiplied as clones and evaluated under comparative yield trials in different situations. Under high density planting in arecanut garden, five hybrids VTLCP-6, VTLCP-20, VTLCP-15, VTLCP-1, and VTLCP-19 have been identified as best performers even in their initial years of growth [28]. Comparative study of parents and progenies as clones resulted in identification of VTLCP-6, VTLCP-2, and VTLCP-20 and parents VTLC-1 and VTLC-56 as potential high yielders [33]. In another trial, clones suitable for both arecanut and coconut canopies have been identified [34] and released as varieties. Under coconut in double hedge system of planting, hybrids VTLCP-22, VTLCP-18, and VTLCP-1 showed the best performance with an optimal canopy and high yield [35]. Evaluation of clonal and seedling progenies of selected genotypes has resulted in identification of four hybrids and two clones for multiplication both as clones and seedlings for utilization in the

To assess the survival and stability of hybrids and clones in different agroclimatic conditions, multi-location trials are important. Elite clones of cocoa are under evaluation in both traditional and nontraditional states, namely Karnataka, Kerala, Tamil Nadu, Andhra Pradesh, Maharashtra, West Bengal, and Assam, for studying genotype location environment interactions. Further, 115 front line

demonstration plots were established under participatory research cum

some degree of homozygosity for the production of hybrids but also to breed materials homozygous for desirable traits like disease resistance. Existence of self-

incompatibility makes inbreeding efforts in cocoa difficult. Since few selfcompatible trees are also identified in the populations, selfing is possible but it should be continued up to six to seven generations to attain homozygosity and thereafter to be utilized for crossing to exploit the hybrid vigor. KAU has taken up this task of selfing self-compatible plants and with over 20 years of continuous effort, maintains two genotypes of S4 generation, 5 of S3, 9 of S2, and 51 genotypes

India is free of most of the debilitating viral and fungal diseases known in cocoa. Since the current cocoa growing area comes under high rainfall zone and the main harvest season coincides with monsoon, incidence of black pod rot caused by Phytophthora palmivora is comparatively higher. Though the main harvest is safe guarded with systematic annual pruning, in the post monsoon period, the second season crop is still affected by pod rot. On field screening, clones have been categorized into having <10, 10–25, and >25% damage levels. In-vitro screening using isolates of P. palmivora, P. capsici, and P. citrophthora indicated that collections of Nigerian origin exhibit certain degree of tolerance [38]. Few Wayanad collections have also expressed field tolerance to pod rot when tested over three seasons. Further, 21 exotic clones collected exclusively for Phytophthora pod rot resistance was identified for utilization in cocoa hybridization program. The variety VTLCH-2, a combination of ICS 6 SCA 6 is found to be tolerant to black pod rot in India as well. KAU has taken up screening and hybridization program for combining desirable traits of CCRP released varieties and black pod resistance in cocoa.

#### 3.6.2 Vascular streak die back

In India, vascular streak die back (VSD) caused by Oncobasidium theobromae was first reported from Kottayam, Kerala, and it began to spread to adjoining cocoa growing areas of the state. As this disease cannot be controlled effectively by the use of fungicides, KAU breeding program concentrated mainly on production of VSD resistant varieties. Hybrid seedlings were screened in the nursery by subjecting them to high inoculum load by keeping them in the midst of infected seedlings. The tolerant and vigorous seedlings were selected and established in field for evaluation. CCRP varieties have been especially released for VSD resistance and also have been utilized for establishment of clonal gardens for seedling supply [31].

#### 3.6.3 Tea mosquito bug

Tea mosquito bug (TMB) (Helopeltis sp.) incidence became severe in the recent years in summer and post monsoon seasons. Helopeltis antonii, H. theivora, and H. bradyi are reported on cocoa in South India. Insect population is influenced by many factors like temperature, humidity, water stress, condition of cocoa tree, etc. The development and use of mirid resistant cocoa varieties is one of the alternatives to chemical control and resistance studies in cocoa have mostly concentrated on assessment of field damage [39]. Damage on flushes, cherelles, and pods of individual trees and different grade levels of infection on cherelles and pods are assessed to work out the TMB tolerance among genotypes. Penetrometer readings for determining the hardness of sclerotic layer, thickness at primary and secondary furrows of pod husk have been recorded in 100 cocoa genotypes and interpreted with reference to insect resistance [40]. Mechanism of plant resistance to insects is a complex phenomenon. Plant attractiveness to some extent affects the level of infestation, antixenosis prevents feeding, while antibiosis disturbs the pest development, and finally cocoa tolerance is assessed from the ability of a tree to contain damage and recover from it. Red colored pods with the smooth surface have been identified as tolerant to TMB damage among Wayanad collections.

be further utilized in the breeding program. Genotypic differences for morphophysiological criteria, potential antioxidant enzymes, and biochemical components depicting drought tolerance in young seedlings were determined with cocoa clones and hybrids under controlled low moisture stress conditions [46–48]. From these trials, standards and thresholds for several physiological parameters related to cocoa

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

Hadley [49] detailed the visual estimates of physiological traits in cocoa, and

DNA fingerprinting with RAPD markers has been done earlier on 76 collections, and the clones VTLC-11, 67, and 83 and VTLC-93 were identified as highly divergent. DNA extraction protocol of cocoa with fully expanded but soft leaves is standardized with the modified SDS method. Recently, 16 SSR primers specific to cocoa were used to assess diversity in 44 exotic clones, and both morphological and molecular diversity were assessed in detail [52]. An attempt has been made to identify the markers for drought sensitivity by utilizing susceptible and tolerant parents and progenies of cocoa [53]. About 75% of the genomic data of cocoa is available in the public domain which has paved the way for analyzing genes related to specific needs. CPCRI hosts one of the Agri Bioinformatics center under the Department of Information Technology and through bioinformatics tools, proteins involved in drought tolerance, Phytophthora resistance, and carotenoid biosynthetic pathways have been analyzed, and databases, CocoaESTdb, CocoaSTRESSdb, and a Standalone EST microsatellite mining and analysis tool (SEMAT) have been

Cocoa improvement has attained a positive phase with the sequencing of its genome. Identifying genes responsible for incompatibility and disease resistance is the main concern of geneticists and molecular biologists. Expression of genes for resistance and quality parameters and their validation with trait specific germplasm is very important for future cocoa improvement program. Possible use of inbred lines will be taken up. Development of early selection, detection, and diagnostic methods for resistance will enable rapid screening of plant material and permit preselection activities. Because of the health benefits of dark chocolates, biochemical

the morpho-physiological parameters include measurements of flowering, fruiting, cherelle wilting, leaf flushing, branching and pruning intensity, canopy shape, density, and light transmission on different point scales. In order to understand and elucidate the optimum canopy shape and structure of cocoa, different spacing and canopy sizes have been studied at CPCRI [50], which showed significant differences in crop yield. In an experiment with grafts, the photosynthetically active radiation (PAR) and light interception varied significantly over the years with two spacings (2.7 2.7 m and 2.7 5.4 m) and three canopy sizes (small, medium, and large), and similar results were noticed with transpiration rate and stomatal conductance [51]. It is important to note that the maximum leaf area should be maintained, self shading of leaves should be avoided, and pruning should be done to the extent of retaining 20–30 leaves/developing pod to ensure the yielding potential of the genotype. With an annual pruning of single tier canopy, fertilizer dose of 100:40:140 g NPK in two splits with 20 L of water as drip is being practiced in maintenance of field gene bank under arecanut and coconut-

were established.

based cropping systems.

developed [54–60].

4. Future prospects

127

3.7 Biotechnology and bioinformatics

DOI: http://dx.doi.org/10.5772/intechopen.82077

#### 3.6.4 Low moisture stress

Cocoa plants are susceptible to environmental conditions especially temperature and drought and considerably influences the pod yield [41]. Cocoa is very sensitive to water scarcity and undergoes a period of low moisture stress for 5–6 months in its current growing condition in India. Detailed study on climate change and weather variability over 43 years (1970–2012) at Vittal, which is located between 12°15<sup>0</sup> N latitude and 75°25<sup>0</sup> E longitude, showed 38% yield variability in cocoa [42]. The trends of temperature increase are +0.4°C for mean maximum (P < 0.001) and +0.4°C for mean minimum during the last decade. Breeding for drought tolerance is unique to our country and is taken up with systematic screening of available germplasm as well as hybridization programs. Screening of accessions is conducted for physiological parameters like stomatal resistance, chlorophyll fluorescence, proline accumulation under stress and by studies on seed germination under low osmotic potential, etc. A total of 216 cocoa genotypes have so far been screened for physiological and biochemical parameters under different trials [43]. In all these studies, field measurements were taken during unstressed (October) and stressed (March) conditions. Few Nigerian collections have been identified as drought tolerant and used for hybridization with high yielding Malaysian collections under two progeny trials. Two hybrids VTLCH-3 and VTLCH-4 have been released as varieties suitable for cultivation under water limited conditions in the country. Studies on leaf morphology, stomatal behavior, water relation components, and biochemical factors indicated that thick leaf, higher wax content, efficient stomatal closure, and high tissue elasticity are responsible for better adaptation of cocoa plants to drought conditions. The application of chlorophyll fluorescence as a tool to screen cocoa for drought tolerance has been confirmed with a series of genotypes. Recently, photosynthesis, chlorophyll fluorescence, and water potential under stress and nonstress conditions were estimated in 11 genotypes from different geographical origins, Columbia, Brazil, Peru, Mexico and Ecuador [44]. Seasonal and varietal differences were found, and transpirational water loss was found to be reduced with increased stomatal closure, which is considered as a favorable drought trait in any crop. Among the 52 new introductions, five Amazon and Pound collections have been found to be adaptable to water limited conditions [45] with high yields, which will

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

be further utilized in the breeding program. Genotypic differences for morphophysiological criteria, potential antioxidant enzymes, and biochemical components depicting drought tolerance in young seedlings were determined with cocoa clones and hybrids under controlled low moisture stress conditions [46–48]. From these trials, standards and thresholds for several physiological parameters related to cocoa were established.

Hadley [49] detailed the visual estimates of physiological traits in cocoa, and the morpho-physiological parameters include measurements of flowering, fruiting, cherelle wilting, leaf flushing, branching and pruning intensity, canopy shape, density, and light transmission on different point scales. In order to understand and elucidate the optimum canopy shape and structure of cocoa, different spacing and canopy sizes have been studied at CPCRI [50], which showed significant differences in crop yield. In an experiment with grafts, the photosynthetically active radiation (PAR) and light interception varied significantly over the years with two spacings (2.7 2.7 m and 2.7 5.4 m) and three canopy sizes (small, medium, and large), and similar results were noticed with transpiration rate and stomatal conductance [51]. It is important to note that the maximum leaf area should be maintained, self shading of leaves should be avoided, and pruning should be done to the extent of retaining 20–30 leaves/developing pod to ensure the yielding potential of the genotype. With an annual pruning of single tier canopy, fertilizer dose of 100:40:140 g NPK in two splits with 20 L of water as drip is being practiced in maintenance of field gene bank under arecanut and coconutbased cropping systems.

### 3.7 Biotechnology and bioinformatics

DNA fingerprinting with RAPD markers has been done earlier on 76 collections, and the clones VTLC-11, 67, and 83 and VTLC-93 were identified as highly divergent. DNA extraction protocol of cocoa with fully expanded but soft leaves is standardized with the modified SDS method. Recently, 16 SSR primers specific to cocoa were used to assess diversity in 44 exotic clones, and both morphological and molecular diversity were assessed in detail [52]. An attempt has been made to identify the markers for drought sensitivity by utilizing susceptible and tolerant parents and progenies of cocoa [53]. About 75% of the genomic data of cocoa is available in the public domain which has paved the way for analyzing genes related to specific needs. CPCRI hosts one of the Agri Bioinformatics center under the Department of Information Technology and through bioinformatics tools, proteins involved in drought tolerance, Phytophthora resistance, and carotenoid biosynthetic pathways have been analyzed, and databases, CocoaESTdb, CocoaSTRESSdb, and a Standalone EST microsatellite mining and analysis tool (SEMAT) have been developed [54–60].

### 4. Future prospects

Cocoa improvement has attained a positive phase with the sequencing of its genome. Identifying genes responsible for incompatibility and disease resistance is the main concern of geneticists and molecular biologists. Expression of genes for resistance and quality parameters and their validation with trait specific germplasm is very important for future cocoa improvement program. Possible use of inbred lines will be taken up. Development of early selection, detection, and diagnostic methods for resistance will enable rapid screening of plant material and permit preselection activities. Because of the health benefits of dark chocolates, biochemical

3.6.3 Tea mosquito bug

3.6.4 Low moisture stress

latitude and 75°25<sup>0</sup>

126

Tea mosquito bug (TMB) (Helopeltis sp.) incidence became severe in the recent

Cocoa plants are susceptible to environmental conditions especially temperature and drought and considerably influences the pod yield [41]. Cocoa is very sensitive to water scarcity and undergoes a period of low moisture stress for 5–6 months in its current growing condition in India. Detailed study on climate change and weather variability over 43 years (1970–2012) at Vittal, which is located between 12°15<sup>0</sup>

trends of temperature increase are +0.4°C for mean maximum (P < 0.001) and +0.4°C for mean minimum during the last decade. Breeding for drought tolerance is unique to our country and is taken up with systematic screening of available germplasm as well as hybridization programs. Screening of accessions is conducted for physiological parameters like stomatal resistance, chlorophyll fluorescence, proline accumulation under stress and by studies on seed germination under low osmotic potential, etc. A total of 216 cocoa genotypes have so far been screened for physiological and biochemical parameters under different trials [43]. In all these studies, field measurements were taken during unstressed (October) and stressed (March) conditions. Few Nigerian collections have been identified as drought tolerant and used for hybridization with high yielding Malaysian collections under two progeny trials. Two hybrids VTLCH-3 and VTLCH-4 have been released as varieties suitable for cultivation under water limited conditions in the country. Studies on leaf morphology, stomatal behavior, water relation components, and biochemical factors indicated that thick leaf, higher wax content, efficient stomatal closure, and high tissue elasticity are responsible for better adaptation of cocoa plants to drought conditions. The application of chlorophyll fluorescence as a tool to screen cocoa for drought tolerance has been confirmed with a series of genotypes. Recently, photosynthesis, chlorophyll fluorescence, and water potential under stress and nonstress conditions were estimated in 11 genotypes from different geographical origins, Columbia, Brazil, Peru, Mexico and Ecuador [44]. Seasonal and varietal differences were found, and transpirational water loss was found to be reduced with increased stomatal closure, which is considered as a favorable drought trait in any crop. Among the 52 new introductions, five Amazon and Pound collections have been found to be adaptable to water limited conditions [45] with high yields, which will

E longitude, showed 38% yield variability in cocoa [42]. The

N

years in summer and post monsoon seasons. Helopeltis antonii, H. theivora, and H. bradyi are reported on cocoa in South India. Insect population is influenced by many factors like temperature, humidity, water stress, condition of cocoa tree, etc. The development and use of mirid resistant cocoa varieties is one of the alternatives to chemical control and resistance studies in cocoa have mostly concentrated on assessment of field damage [39]. Damage on flushes, cherelles, and pods of individual trees and different grade levels of infection on cherelles and pods are assessed to work out the TMB tolerance among genotypes. Penetrometer readings for determining the hardness of sclerotic layer, thickness at primary and secondary furrows of pod husk have been recorded in 100 cocoa genotypes and interpreted with reference to insect resistance [40]. Mechanism of plant resistance to insects is a complex phenomenon. Plant attractiveness to some extent affects the level of infestation, antixenosis prevents feeding, while antibiosis disturbs the pest development, and finally cocoa tolerance is assessed from the ability of a tree to contain damage and recover from it. Red colored pods with the smooth surface have been

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

identified as tolerant to TMB damage among Wayanad collections.

constituents and antioxidant properties of cocoa are to be given greater attention in the breeding programs. Farmers participatory plant breeding, in-situ conservation of land races, exploitation of flavor components from genotypes belonging to specific geographic region, varieties for changing climatic conditions, and environment-friendly management strategies will be considered. Adaptability of cocoa genotypes in traditional and nontraditional zones should be verified, and location specific varieties should be developed [61]. At the national level, expansion of cocoa cultivation with the quality planting material of elite clones, collaborative approach between research institutes, universities, state horticulture departments, and developmental agencies are required. At the international level, participation of India in cocoa genetic resources networking and regional breeding groups of both developed and developing countries is important. Collaboration of India with Asia Pacific regional countries, Malaysia, Indonesia, Philippines, Vietnam, and Papua New Guinea is essential with their common coconut-based cropping systems with known pest and disease problems. This will enhance region specific sustainability of cocoa cultivation.

References

1964. p. 27

[1] Ratnam R. Introduction of Criollo cocoa into Madras state. South Indian

DOI: http://dx.doi.org/10.5772/intechopen.82077

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

Food and Agricultural Organisation of the United Nations, International Board for Plant Genetic Resources; 1989

[10] Michelle EJ, Daymond AJ, Hadley P. Technical Guidelines for the Safe Movement of Cacao Germplasm. Revised from FAO/IPGRI Technical Guidelines no. 20. Global Cacao Genetic

Resources Network (CacaoNet). Montpellier, France: Bioversity International; 2014. p. 69

[11] International Cocoa Germplasm Database. UK: University of Reading. Available from: www.icgd.rdg.ac.uk

[12] Bartley BGD. The Genetic Diversity of Cocoa and its Utilization. UK: CABI

[13] Wood GAR, Lass RA. Cocoa. 4th ed. England: Longman Group Limited;

[14] Turnbull CS, Eskes AB. A Visual Aid to Identify Widely Distributed Cocoa

[15] Elain AS, Rajan P. Profile of Cocoa Collections at CPCRI, Research Centre, Kannara. Kasaragod: CPCRI; 2009. p. 61

Arunachalam V, editors. Improvement of Plantation Crops. Kasaragod: CPCRI;

[16] Bhat VR. Cocoa germplasm characterization, conservation and utilization. In: Ratnambal MJ, Kumaran

PM, Muralidharan K, Niral V,

[17] Bekele FL, End MJ, Eskes AB. Proceedings of the International Workshop on Cocoa Breeding for Improved Production Systems; 19–21st October 2003. Accra, Ghana: INGENIC;

1999. pp. 77-85

2005. p. 198

Accessions. Amsterdam, The Netherlands: CFC, UK: ICCO, Italy: Bioversity International; 2010. p. 43

publishing; 2005. p. 341

1985. p. 620

[2] Wood GAR. Cocoa Growing in India. Bournville, UK: Cadbury Brothers Limited, Publication Department;

[3] Directorate of Cashewnut and Cocoa Development, Ministry of Agriculture and Farmers Welfare, Govt. of India. Available from: www.dccd.nic.in

PrasannakumariAmma S. Technologies for increasing productivity in cocoa. In: Bhat R, Balasimha D, Jayasekhar S, editors. Proceedings of National

Seminar on Technologies for Enhancing Productivity in Cocoa. Regional Station,

Balasimha D. Cocoa breeding at CPCRI, India. INGENIC Newsletter. 2005;10:

[7] Elain Apshara S. Cocoa improvement programmes of ICAR—CPCRI—A glance. The Cashew and Cocoa Journal.

[8] Hadley P, Lee T. Current cocoa quarantine facilities of the University of Reading. In: International Workshop on Conservation, Characterization and Utilization of Cocoa Genetic Resources in the 21st Century, Trinidad; 1317 September 1992. 1992. pp. 65-68

[9] Frison EA, Feliu E. FAO/IBPGR Technical Guidelines for the Safe Movement of Cacao Germplasm. Rome:

[4] Bekele FL. Proceedings of the International Workshop on New Technologies and Cocoa Breeding; 16–17th October 2000. Kota Kinabalu, Sabah, Malaysia: INGENIC; 2001. p. 190

[5] Peter KV, Mallika VK,

Vittal: CPCRI; 2002. pp. 1-11

34-37

129

2017;6(2):9-19

[6] Elain Apshara S, Ananda KS,

Horticulture. 1961;9:24-29

## Author details

Subbian Elain Apshara ICAR—Central Plantation Crops Research Institute-CPCRI, Vittal, Karnataka, India

\*Address all correspondence to: elain\_apshara@yahoo.co.in

© 2019 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited.

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

## References

constituents and antioxidant properties of cocoa are to be given greater attention in the breeding programs. Farmers participatory plant breeding, in-situ conservation of land races, exploitation of flavor components from genotypes belonging to spe-

environment-friendly management strategies will be considered. Adaptability of cocoa genotypes in traditional and nontraditional zones should be verified, and location specific varieties should be developed [61]. At the national level, expansion of cocoa cultivation with the quality planting material of elite clones, collaborative approach between research institutes, universities, state horticulture departments, and developmental agencies are required. At the international level, participation of India in cocoa genetic resources networking and regional breeding groups of both developed and developing countries is important. Collaboration of India with Asia Pacific regional countries, Malaysia, Indonesia, Philippines, Vietnam, and Papua New Guinea is essential with their common coconut-based cropping systems with known pest and disease problems. This will enhance region specific sustainability of

ICAR—Central Plantation Crops Research Institute-CPCRI, Vittal, Karnataka, India

© 2019 The Author(s). Licensee IntechOpen. This chapter is distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/ by/3.0), which permits unrestricted use, distribution, and reproduction in any medium,

\*Address all correspondence to: elain\_apshara@yahoo.co.in

provided the original work is properly cited.

cific geographic region, varieties for changing climatic conditions, and

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

cocoa cultivation.

Author details

128

Subbian Elain Apshara

[1] Ratnam R. Introduction of Criollo cocoa into Madras state. South Indian Horticulture. 1961;9:24-29

[2] Wood GAR. Cocoa Growing in India. Bournville, UK: Cadbury Brothers Limited, Publication Department; 1964. p. 27

[3] Directorate of Cashewnut and Cocoa Development, Ministry of Agriculture and Farmers Welfare, Govt. of India. Available from: www.dccd.nic.in

[4] Bekele FL. Proceedings of the International Workshop on New Technologies and Cocoa Breeding; 16–17th October 2000. Kota Kinabalu, Sabah, Malaysia: INGENIC; 2001. p. 190

[5] Peter KV, Mallika VK, PrasannakumariAmma S. Technologies for increasing productivity in cocoa. In: Bhat R, Balasimha D, Jayasekhar S, editors. Proceedings of National Seminar on Technologies for Enhancing Productivity in Cocoa. Regional Station, Vittal: CPCRI; 2002. pp. 1-11

[6] Elain Apshara S, Ananda KS, Balasimha D. Cocoa breeding at CPCRI, India. INGENIC Newsletter. 2005;10: 34-37

[7] Elain Apshara S. Cocoa improvement programmes of ICAR—CPCRI—A glance. The Cashew and Cocoa Journal. 2017;6(2):9-19

[8] Hadley P, Lee T. Current cocoa quarantine facilities of the University of Reading. In: International Workshop on Conservation, Characterization and Utilization of Cocoa Genetic Resources in the 21st Century, Trinidad; 1317 September 1992. 1992. pp. 65-68

[9] Frison EA, Feliu E. FAO/IBPGR Technical Guidelines for the Safe Movement of Cacao Germplasm. Rome: Food and Agricultural Organisation of the United Nations, International Board for Plant Genetic Resources; 1989

[10] Michelle EJ, Daymond AJ, Hadley P. Technical Guidelines for the Safe Movement of Cacao Germplasm. Revised from FAO/IPGRI Technical Guidelines no. 20. Global Cacao Genetic Resources Network (CacaoNet). Montpellier, France: Bioversity International; 2014. p. 69

[11] International Cocoa Germplasm Database. UK: University of Reading. Available from: www.icgd.rdg.ac.uk

[12] Bartley BGD. The Genetic Diversity of Cocoa and its Utilization. UK: CABI publishing; 2005. p. 341

[13] Wood GAR, Lass RA. Cocoa. 4th ed. England: Longman Group Limited; 1985. p. 620

[14] Turnbull CS, Eskes AB. A Visual Aid to Identify Widely Distributed Cocoa Accessions. Amsterdam, The Netherlands: CFC, UK: ICCO, Italy: Bioversity International; 2010. p. 43

[15] Elain AS, Rajan P. Profile of Cocoa Collections at CPCRI, Research Centre, Kannara. Kasaragod: CPCRI; 2009. p. 61

[16] Bhat VR. Cocoa germplasm characterization, conservation and utilization. In: Ratnambal MJ, Kumaran PM, Muralidharan K, Niral V, Arunachalam V, editors. Improvement of Plantation Crops. Kasaragod: CPCRI; 1999. pp. 77-85

[17] Bekele FL, End MJ, Eskes AB. Proceedings of the International Workshop on Cocoa Breeding for Improved Production Systems; 19–21st October 2003. Accra, Ghana: INGENIC; 2005. p. 198

[18] Eskes AB, Lanaud C. Cocoa. In: Charrier A, Jacquot M, Hamon S, Nicolas D, editors. Tropical Plant Breeding. USA: Science Publishers Inc., France: CIRAD; 2001. pp. 78-105

[19] Elain Apshara S, Bhat VR, Ananda KS, Nair RV, Suma D. Evaluation and identification of high yielding trees in Nigerian cocoa germplasm. Journal of Plantation Crops. 2009;37(2):111-116

[20] Elain Apshara S, Nair RV. Genetic analysis in cocoa collections obtained from Nigeria. Journal of Plantation Crops. 2011;39(1):200-206

[21] Elain Apshara S. Growth and yield performance of Trinidad cocoa collections in Karnataka. Journal of Plantation Crops. 2015;43(3):237-240

[22] Elain Apshara S. Selection of potential clones from Wayanad cocoa collections. Journal of Plantation Crops. 2016;44(2):124-128

[23] GOI. Cocoa beans grading and marking rules. AGMARK standards. Gazette of India Part II. 1997;3:1

[24] Minimol JS, PrasannakumariAmma S, Lalitha Bai EK. Three decades of research on breeding of cocoa in Kerala Agricultural University. In: Elain Apshara S, Jaganathan D, Balasimha D, editors. Proceedings of Seminar on Strategies for Enhancing Productivity of Cocoa. Regional Station, Vittal: CPCRI; 2011. pp. 10-14

[25] Senthil Amudhan M, Elain Apshara S. A comparative study on antioxidant activity and biochemical profile of exotic cocoa clones. Journal of Plantation Crops. 2015;43(3):231-235

[26] Vikraman NR, Mallika VK, PrasannakumariAmma S. Breeding and genetics. In: Balasimha D, editor. Cocoa. Kasaragod: CPCRI; 2002. pp. 18-47

[27] Elain Apshara S, Bhat VR, Ananda KS, Nair RV. High yielding cocoa varieties of the Central Plantation Crops Research Institute, India. INGENIC Newsletter. 2007;11:12-15

clones under arecanut and coconut. Journal of Plantation Crops. 2013;41(2):

DOI: http://dx.doi.org/10.5772/intechopen.82077

[41] Daymond AJ, Hadley P, Machado RCR, Ng E. An investigation of physiological parameters underlying yield variation in cocoa. In: Proceedings of 13th International Cocoa Research Conference; 9–14th October, 2000. Vol. I. Kota Kinabalu, Sabah, Malaysia;

[42] Sujatha S, Bhat R, Elain Apshara S. Climate change, weather variability and associated impact on arecanut and cocoa in humid tropics of India. International Journal of Innovative Horticulture.

[43] Balasimha D. Stress physiology of cocoa. Journal of Plantation Crops. 1999;

tolerance. Journal of Plantation Crops.

[45] Balasimha D, Elain Apshara S, Jose CT. Genotypic variations in chlorophyll fluorescence and stomatal conductance of cocoa in relation to drought tolerance. Journal of Plantation Crops. 2013;41(1):

[46] Antoine Alban M'b K, Elain Apshara S, Hebbar KB, Mathias TG, Sévérin A. Potential of antioxidant enzymes in depicting drought tolerance in cocoa genotypes at young age.

African Journal of Science and Research.

[47] Alban M'b KA, Elain Apshara S, Hebbar KB, Mathias TG, Sévérin A. Morpho-physiological criteria for assessment of two month old cocoa genotypes for drought tolerance. Indian Journal of Plant Physiology. 2015a;21(1):

[44] Elain Apshara S, Rajesh MK, Balasimha D. Assessment of morphological, physiological and molecular characteristics of cocoa accessions from Central and South America in relation to drought

2000. pp. 331-340

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

2017;7(1):27-37

2013;41(3):389-397

40-45

23-30

2015;4(5):18-23

27(1):1-8

[35] Elain Apshara S. Performance of elite cocoa clones under coconut in double hedge system of planting. The Cashew and Cocoa Journal. 2014;3(2):

[36] Elain Apshara S. Comparative study on clonal and seedling progenies of selected cocoa genotypes. Indian Journal of Horticulture. 2017;74(2):168-172

[37] Daniel M. Pests management of cocoa. In: Bhat R, Balasimha D, Jayasekhar S, editors. Proceedings of National Seminar on Technologies for Enhancing Productivity in Cocoa. Regional Station, Vittal: CPCRI; 2002.

[38] Chandramohanan R. Studies on the reaction of pods of different cocoa accessions to Phytophthora palmivora.

[39] N'Guessan KF, N'Goran JAK, Eskes AB. Mirid resistance studies in Cote d'Ivoire: Assessment of antixenosis, antibiosis and tolerance. In: Eskes AB, Efron Y, editors. Global Approaches to Cocoa Germplasm Utilization and Conservation. Final report of the CFC/ICCO/IPGRI project on Cocoa

242-246

13-16

pp. 87-94

Planter. 1982;58:99-103

Germplasm Utilization and Conservation (1998–2004). CFC Technical Paper No.50. CFC,

pp. 177-186

131

Amsterdam, The Netherlands/ICCO, London, UK/ IPGRI, Rome, Italy. 2004.

[40] Elain Apshara S. Selecting elite cocoa clones for phenotypic characters including pod wall hardness in India. In: Samsudin S, Chee SYK, Zainudin BH, editors. Abstracts of Malaysian International Cocoa Conference on Innovation and Technology Driving Cocoa Productivity/Sustainability; 7–8 October 2013. Kuala Lumpur: Sunway Pyramid Convention Centre; 2013. p. 69

[28] Elain Apshara S, Bhat VR, Nair RV. Comparative studies on elite cocoa progenies in their initial years of growth. Journal of Plantation Crops. 2008;36(1):38-44

[29] Elain Apshara S. Cocoa Varieties of ICAR-CPCRI. Extension Folder No. 249, Centenary Publication No. 45. Kasaragod: CPCRI; 2016

[30] Elain AS. Cocoa Planting Material Production. Technical Bulletin No. 66. Kasaragod: CPCRI; 2010. p. 24

[31] Mallika VK, Prasannakumari Amma S, Abraham K, Vikraman Nair R, Minimol JS. Evolution of cocoa varieties resistant to vascular streak die back through hybridization. In: Abstr. Inter. Conf. Plantation Crops. PLACROSYM XIV; 12–15 December 2000. Hyderabad; 2000. p. 6

[32] Mallika VK, PrasannakumariAmma S, Vikraman Nair R. Crop improvement in cocoa. In: Bhat R, Balasimha D, Jayasekhar S, editors. Proceedings of National Seminar on Technologies for Enhancing Productivity in Cocoa. Regional Station, Vittal: CPCRI; 2002. pp. 19-27

[33] Elain Apshara S. Comparative performance of selected cocoa parents and progenies as clones in South India. In: Samsudin S, Chee SYK, Zainudin BH, editors. Proceedings of Malaysian International Cocoa Conference on Innovation and Technology Driving Cocoa Productivity/Sustainability; 7–8 October 2013. Kuala Lumpur: Sunway Pyramid Convention Centre; 2013. pp. 9-17

[34] Elain Apshara S. Performance of selected cocoa (Theobroma cacao L.)

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

clones under arecanut and coconut. Journal of Plantation Crops. 2013;41(2): 242-246

[18] Eskes AB, Lanaud C. Cocoa. In: Charrier A, Jacquot M, Hamon S, Nicolas D, editors. Tropical Plant Breeding. USA: Science

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

[27] Elain Apshara S, Bhat VR, Ananda KS, Nair RV. High yielding cocoa varieties of the Central Plantation Crops Research Institute, India. INGENIC

[28] Elain Apshara S, Bhat VR, Nair RV. Comparative studies on elite cocoa progenies in their initial years of growth. Journal of Plantation Crops.

[29] Elain Apshara S. Cocoa Varieties of ICAR-CPCRI. Extension Folder No. 249,

[30] Elain AS. Cocoa Planting Material Production. Technical Bulletin No. 66.

[31] Mallika VK, Prasannakumari Amma S, Abraham K, Vikraman Nair R, Minimol JS. Evolution of cocoa varieties resistant to vascular streak die back through hybridization. In: Abstr. Inter. Conf. Plantation Crops. PLACROSYM XIV; 12–15 December 2000. Hyderabad;

[32] Mallika VK, PrasannakumariAmma S, Vikraman Nair R. Crop improvement in cocoa. In: Bhat R, Balasimha D, Jayasekhar S, editors. Proceedings of National Seminar on Technologies for Enhancing Productivity in Cocoa. Regional Station, Vittal: CPCRI; 2002.

[33] Elain Apshara S. Comparative performance of selected cocoa parents and progenies as clones in South India. In: Samsudin S, Chee SYK, Zainudin BH, editors. Proceedings of Malaysian International Cocoa Conference on Innovation and Technology Driving Cocoa Productivity/Sustainability; 7–8 October 2013. Kuala Lumpur: Sunway Pyramid Convention Centre; 2013.

[34] Elain Apshara S. Performance of selected cocoa (Theobroma cacao L.)

Centenary Publication No. 45. Kasaragod: CPCRI; 2016

Kasaragod: CPCRI; 2010. p. 24

Newsletter. 2007;11:12-15

2008;36(1):38-44

2000. p. 6

pp. 19-27

pp. 9-17

Publishers Inc., France: CIRAD; 2001.

[19] Elain Apshara S, Bhat VR, Ananda KS, Nair RV, Suma D. Evaluation and identification of high yielding trees in Nigerian cocoa germplasm. Journal of Plantation Crops. 2009;37(2):111-116

[20] Elain Apshara S, Nair RV. Genetic analysis in cocoa collections obtained from Nigeria. Journal of Plantation

[21] Elain Apshara S. Growth and yield

Crops. 2011;39(1):200-206

performance of Trinidad cocoa collections in Karnataka. Journal of Plantation Crops. 2015;43(3):237-240

[22] Elain Apshara S. Selection of potential clones from Wayanad cocoa collections. Journal of Plantation Crops.

[23] GOI. Cocoa beans grading and marking rules. AGMARK standards. Gazette of India Part II. 1997;3:1

[24] Minimol JS, PrasannakumariAmma S, Lalitha Bai EK. Three decades of research on breeding of cocoa in Kerala Agricultural University. In: Elain Apshara S, Jaganathan D, Balasimha D, editors. Proceedings of Seminar on Strategies for Enhancing Productivity of Cocoa. Regional Station, Vittal: CPCRI;

[25] Senthil Amudhan M, Elain Apshara S. A comparative study on antioxidant activity and biochemical profile of exotic cocoa clones. Journal of Plantation Crops. 2015;43(3):231-235

PrasannakumariAmma S. Breeding and genetics. In: Balasimha D, editor. Cocoa. Kasaragod: CPCRI; 2002. pp. 18-47

[26] Vikraman NR, Mallika VK,

2016;44(2):124-128

2011. pp. 10-14

130

pp. 78-105

[35] Elain Apshara S. Performance of elite cocoa clones under coconut in double hedge system of planting. The Cashew and Cocoa Journal. 2014;3(2): 13-16

[36] Elain Apshara S. Comparative study on clonal and seedling progenies of selected cocoa genotypes. Indian Journal of Horticulture. 2017;74(2):168-172

[37] Daniel M. Pests management of cocoa. In: Bhat R, Balasimha D, Jayasekhar S, editors. Proceedings of National Seminar on Technologies for Enhancing Productivity in Cocoa. Regional Station, Vittal: CPCRI; 2002. pp. 87-94

[38] Chandramohanan R. Studies on the reaction of pods of different cocoa accessions to Phytophthora palmivora. Planter. 1982;58:99-103

[39] N'Guessan KF, N'Goran JAK, Eskes AB. Mirid resistance studies in Cote d'Ivoire: Assessment of antixenosis, antibiosis and tolerance. In: Eskes AB, Efron Y, editors. Global Approaches to Cocoa Germplasm Utilization and Conservation. Final report of the CFC/ICCO/IPGRI project on Cocoa Germplasm Utilization and Conservation (1998–2004). CFC Technical Paper No.50. CFC, Amsterdam, The Netherlands/ICCO, London, UK/ IPGRI, Rome, Italy. 2004. pp. 177-186

[40] Elain Apshara S. Selecting elite cocoa clones for phenotypic characters including pod wall hardness in India. In: Samsudin S, Chee SYK, Zainudin BH, editors. Abstracts of Malaysian International Cocoa Conference on Innovation and Technology Driving Cocoa Productivity/Sustainability; 7–8 October 2013. Kuala Lumpur: Sunway Pyramid Convention Centre; 2013. p. 69 [41] Daymond AJ, Hadley P, Machado RCR, Ng E. An investigation of physiological parameters underlying yield variation in cocoa. In: Proceedings of 13th International Cocoa Research Conference; 9–14th October, 2000. Vol. I. Kota Kinabalu, Sabah, Malaysia; 2000. pp. 331-340

[42] Sujatha S, Bhat R, Elain Apshara S. Climate change, weather variability and associated impact on arecanut and cocoa in humid tropics of India. International Journal of Innovative Horticulture. 2017;7(1):27-37

[43] Balasimha D. Stress physiology of cocoa. Journal of Plantation Crops. 1999; 27(1):1-8

[44] Elain Apshara S, Rajesh MK, Balasimha D. Assessment of morphological, physiological and molecular characteristics of cocoa accessions from Central and South America in relation to drought tolerance. Journal of Plantation Crops. 2013;41(3):389-397

[45] Balasimha D, Elain Apshara S, Jose CT. Genotypic variations in chlorophyll fluorescence and stomatal conductance of cocoa in relation to drought tolerance. Journal of Plantation Crops. 2013;41(1): 40-45

[46] Antoine Alban M'b K, Elain Apshara S, Hebbar KB, Mathias TG, Sévérin A. Potential of antioxidant enzymes in depicting drought tolerance in cocoa genotypes at young age. African Journal of Science and Research. 2015;4(5):18-23

[47] Alban M'b KA, Elain Apshara S, Hebbar KB, Mathias TG, Sévérin A. Morpho-physiological criteria for assessment of two month old cocoa genotypes for drought tolerance. Indian Journal of Plant Physiology. 2015a;21(1): 23-30

[48] Antoine Alban M'b K, Elain Apshara S, Hebbar KB, Ananda KS, Mathias TG, Sévérin A. Change in epicuticular wax and biochemical secondary metabolites in cocoa under hydric stress. International Journal of Current Research. 2016;8(4): 28988-28999

[49] Hadley P. Physiological traits. In: Eskes AB, Engels JMM, Lass RA, editors. Working Procedure for Cocoa Germplasm Evaluation and Selection. Rome, Italy: IPGRI; 2000. pp. 91-94

[50] Balasimha D. Growth and yield of cocoa grown under areca in relation to plant density and canopy architecture. In: Proceedings of 13th International Cocoa Research Conference. Kota Kinabalu, Malaysia; 2001. pp. 365-372

[51] Balasimha D. Performance of cocoa in relation to spacing and pruning regimes. In: Proceedings of 15th International Cocoa Research Conference; 9–4 October 2006. Costa Rica; 2006. pp. 265-272

[52] Elain Apshara S, Rajesh MK. Evaluation of exotic cocoa collections in India for their morphological and molecular diversity. In: International Symposium on Frontiers in Science and Technology for Cacao Quality, Productivity and Sustainability; 31 May–3 June 2016. USA: Penn State College of Agricultural Sciences, University Park; 2016. Book of abstracts: 44

[53] Alban M'b KA, Elain Apshara S, Rahman S, Rajesh MK, Mathias TG, Sévérin A. Analysis of drought tolerance in cocoa genotypes and their hybrids by physiological parameters and molecular markers. In: Abstracts of PLACROSYM XXI; 10–12 December. Calicut: IISR; 2014. p. 43

[54] Naganeeswaran S, Manimekalai R, Elain Apshara S, Manju KP, Malhotra SK, Karun A. Standalone EST Microsatellite Mining and Analysis Tool (SEMAT) for automated EST-SSR analysis in plants Tree Genetics and Genomics. 2014;10(6):1755-1757

[60] Elain Apshara S, Naganeeswaran S,

DOI: http://dx.doi.org/10.5772/intechopen.82077

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India

Manimekalai R. CocoaESTdb: A comprehensive cocoa Expressed Sequence Tag database. In: Abstracts of National Seminar on Applications of Bioinformatics in Agriculture; 11–12 November 2013. Kasaragod: CPCRI;

[61] Malhotra SK, Elain Apshara S. Genetic resources of cocoa and their utilization—An appraisal. Indian Journal

of Genetics. 2017;77(2):199-213

2013c. p. 51

133

[55] Naganeeswaran S, Elain Apshara S, Manimekalai R. Analysis of expressed sequence tags (ESTs) from cocoa (Theobroma cacao L) upon infection with Phytophthora megakarya. Bioinformation. 2012;8(2):65-69

[56] Naganeeswaran S, Elain Apshara S, Manimekalai R. Bioinformatics tools for EST/transcriptome data analysis. In: Abstracts of National Seminar on Applications of Bioinformatics in Agriculture; 11–12 November 2013. Kasaragod: CPCRI; 2013. p. 53

[57] Naganeeswaran S, Elain Apshara S, Manimekalai R, Alban M'BKA. Cocoa stress gene database (CocoaSTRESSdb): Comprehensive resource for cocoa stress response genes and associated SSR markers to facilitate marker assisted selection. In: Abstracts of National Seminar on New Horizons and Challenges in Biotechnology and Bioinformatics; 09–10 October 2014. Kasaragod: CPCRI; 2014. p. 65

[58] Naganeeswaran S, Manimekalai R, Elain Apshara S, Manju KP, Malhotra SK, Karun A. Standalone EST microsatellite mining and analysis tool (SEMAT) for automated EST-SSR analysis in plants. Tree Genetics and Genomics. 2014. DOI: 10.1007/ s11295-014-0785-2

[59] Naganeeswaran S, Elain Apshara S, Manimekalai R, Vasu A, Malhotra SK. Cocoa EST database: Comprehensive database of cocoa expressed sequence tags (ESTs). International Journal of Innovative Research in Computer and Communication Engineering. 2015; 3(11):10441-10444

Cocoa Genetic Resources and Their Utilization in Palm-Based Cropping Systems of India DOI: http://dx.doi.org/10.5772/intechopen.82077

[60] Elain Apshara S, Naganeeswaran S, Manimekalai R. CocoaESTdb: A comprehensive cocoa Expressed Sequence Tag database. In: Abstracts of National Seminar on Applications of Bioinformatics in Agriculture; 11–12 November 2013. Kasaragod: CPCRI; 2013c. p. 51

[48] Antoine Alban M'b K, Elain Apshara S, Hebbar KB, Ananda KS, Mathias TG, Sévérin A. Change in epicuticular wax and biochemical secondary metabolites in cocoa under hydric stress. International Journal of

Theobroma cacao - Deploying Science for Sustainability of Global Cocoa Economy

[54] Naganeeswaran S, Manimekalai R, Elain Apshara S, Manju KP, Malhotra

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