Domestication and breeding of peach palm

1 Biologist, M.S., PhD., Researcher at the National Research Institute for Amazonia (INPA)and CNPq Fellowship. Email: [email protected]

2 Agronomist, M.S., D.S. and Researcher at Embrapa Florestas. Email: [email protected] Agronomist, M.S., D.S. and Researcher at the Instituto Agronômico. Email:

[email protected] Agronomist, M.S., D.S., Researcher at the Instituto Nacional de Pesquisas da Amazônia

(INPA) and CNPq Fellowship. Email: [email protected] Biologist, M.S., D.S. and Professor at the Federal University of Amazonas. Email:

[email protected] Forestry Engineer, M.S. and Researcher at the National Research Institute for Amazonia

(INPA). Email: [email protected] Agronomist, M.S., D.S. and Researcher at Embrapa Amazônia Oriental. Email:

[email protected] Biologist, M.S. and National Research Institute for Amazonia (INPA) Scholarship. Email:

[email protected] Agronomist, M.S., D.S. and Foundation for the Support of Research in the State of

Amazonas (FAPEAM) Scholarship. Email: [email protected]

chapter 18

Domestication and Breeding of thePeach Palm

Charles R. Clement1, Antônio Nascim Kalil Filho2, Valéria A. Modolo3

Kaoru Yuyama4, Doriane Picanço Rodrigues5, Johannes van Leeuwen6

João Tomé de Farias Neto7

Introduction

The peach palm or pupunha (Bactris gasipaes Kunth, Palmae) is theonly domesticated palm in the Neotropics (Clement, 1988), although other palmshave been semi-domesticated or are in the early stages of domestication (Clement,1999). It is believed that the Amerindians became interested in the species for itswood, preferred for making tools for hunting and fishing, as well as forconstruction (Patiño, 1989). Over a period of thousands of years, its starchy-

Clement et al.362

oily fruit became more important. Before the European conquest, harvesting thisstarchy fruit was cause for celebration among the peoples of north-westernAmazonia, the Pacific coast of Colombia and southern Central America (Patiño,2002). After the conquest, the peach palm gradually lost its importance, asconsumer preferences changed with the introduction of exotic crops, but itcontinues to be reasonably popular wherever it is found.

Throughout the colonial period, we find constant references to thequalities of the peach palm, always in connection with its importance toindigenous peoples and indicating that it is a fruit that improves with cooking.However, it attracted little attention until the beginning of the 20

th century.

Popenoe and Jimenez (1921) aroused modern interest in the peach palm, not justin Central America but also in Brazil. Over the next few decades, variousinstitutions in a number of countries began studies that gradually providedinformation on its nutritional properties, propagation, cultivation, uses andpreferences of consumers in a variety of locations (Mora Urpí et al., 1997).However, interest in the crop did not increase significantly.

In the second half of the 20th century, interest was aroused by a completely

different product of the peach palm: hearts-of-palm. The first experiment in palmheart production was carried out at the end of the 1960s (Camacho and Soria,1970), changing the future of the peach palm. Over the next decade, it wasplanted for its hearts-of-palm in Costa Rica and later in Brazil, Panama, Ecuador,Colombia, Peru, Venezuela and Nicaragua, becoming a profitable crop in manycountries (Mora Urpí et al., 1997). The expansion of palm heart plantations leadto the creation of new breeding programs in Brazil and other countries, especiallyCosta Rica. At present, Brazil is the largest producer, followed by Ecuador andCosta Rica, with Ecuador and Costa Rica the leading exporters to the internationalmarket, since Brazil consumes almost all the peach palm products it produces(Clement, 2008).

Note that the peach palm is cultivated in two distinct ways: for its fruitand for its hearts-of-palm. It is cultivated for its fruit in family orchards, smallagroforestry systems and occasionally in small monoculture orchards at densitiesof a few hundred palms per hectare. The plants reach reproductive phase. It iscultivated for palm hearts in monoculture plantations with five to ten thousandpalms per hectare. In this type of cropping, the plant is cut to obtain the palmheart shortly after the stem appears at soil level, long before it reaches the

363Domestication and beeding of the peach palm

reproductive phase. If the clump is well looked after, it can survive decades ofcutting without ever reaching the reproductive phase.

This chapter up-dates the state-of-the-art for topics directly relevant tounderstanding the origin, domestication and breeding of the peach palm, addingnew information to the revision of Mora Urpí et al. (1997). All the informationgiven with no citation can be sourced in that revision, which also containsinformation on the agronomy and uses of the peach palm. Next, this chaptercovers the main peach palm breeding projects for both fruit and heart-of-palmproduction in Brazil, showing the variety of ideas and practices applied. Finally,we discuss the future prospects for peach palm and related breeding projects.

Taxonomy

The peach palm is a multi-stemmed palm that can reach a height of 20 m(Figure 18.1). Its fibrous root system extends up to 7 m from the stem, penetratingto a depth of 2 m. The stem diameter ranges from 15 to 30 cm and internodelength from 1 to 30 cm (the node is the scar left when a senescent leaf falls away).Internodes bear numerous stiff black or dark brown spines, but there are spinelessmutations that have been selected by the Amerindians in various regions. Theapex of the stem supports a crown of 15 to 25 pinnate leaves, with leafletsinserted at different angles. The tender, unexpanded leaves above the meristemin the middle of the crown form the heart-of-palm. The monoecious inflorescenceappears in the axillae of senescing leaves. After pollination, bunches can containbetween 50 and 1,000 fruits and weigh from 1 to 25 kg. Various factors, such asplant nutrition, deficient pollination, drought, competition and attack by pestsand diseases, can cause the fruits to abort, reducing the bunch weight or evenaborting the whole bunch. The fruit weighs between 0.5 and 250 g, and istraditionally the main economic product. When mature, it has a fibrous epicarpthat varies in color from red to orange to yellow, a mesocarp that varies fromstarchy to oily, and an endocarp surrounding a fibrous and oily kernel.

The peach palm fruit is extremely variable in terms of shape, size, exocarpand mesocarp (due to different beta-carotene levels) (Figure 18.2), chemicalcomposition (Table 18.1), texture, flavor and aroma. This enormous variabilityexplains why its taxonomic history is so confused.

Clement et al.364

Figure 18.1. Peach palm plantation at the onset of flowering, spaced at 5 x 5 m.

Figure 18.2. Peach palm fruit bunches of the Pará landrace (yellow fruits; ~ 20 g),Solimões landrace (red fruits; ~ 30 g; and orange fruits with “rayas”(brown cracks in the exocarp); ~ 25 g) and variety chichagui type 3(small orange-red fruits; ~ 2 g). Color is not a landrace characteristic.


Table 18.1. Fruit fresh weight, percentage of fresh mesocarp, chemical compositionof the mesocarp and productivity of plants in two wild peach palmpopulations (Bactris gasipaes var. chichagui, type 1 – Acre andtype 3 – Ucayali) and four landraces (B. gasipaes var. gasipaes)along the domestication continuum (Clement, 2006).

Fru

it

Pu

lp

Wa

ter

Pro

tein

Oil

Ca

rbo

.

Fib

er

En

erg

y

Pro

d.

Name Locality g % g / 100 g kcal kg

Acre Rio Branco, AC

1 60 41,1 5,4 27,1 0,3 24,6 365 2

Ucayali Pucallpa, Peru

1,4 66 42,8 3,4 27,2 12 13,8 362 3

Juruá Cruzeiro do Sul, AC

20 80 54,4 3,1 13,8 19,6 8,4 249 12

Solimões Coarí, AM 58 91 42,7 4,1 12 31,2 9,3 286 36

Putumayo Benj. Constant, AM

70 96 52,6 1,9 3,5 38 3,2 204 30

Vaupés São Gabriel, AM

83 96 60,9 2,4 2,6 31,6 1,8 167 30

Fonte: Clement (2006)

The first name given to the peach palm was Martinezia ciliata Ruiz andPavon (1798), based on various wild populations growing in the Peruvian Andes,south of Yurimaguas. Two decades later, Kunth (1816) gave the name Bactrisgasipaes to the cultivated populations observed by Humboldt and Bonplandalong the Magdalena River in Colombia. Shortly afterwards, Martius (1824) namedit Guilielma speciosa, based on cultivated populations observed in easternPará state, Brazil. Throughout the 19

th and 20

th centuries, nine other species and

five varieties were described, some in Bactris and others in Guilielma. Theconfusion was so great that the peach palm was considered a cultigen, acultivated species with no known wild relative (Schultes, 1984).

In 2000, Henderson published a revision of Bactris, creating two varietiesto combine previous species and varieties: B. gasipaes var. gasipaes, whichincludes all domesticated peach palm populations with large fruits (10 to 250 g);and B. gasipaes var. chichagui (H. Karsten) Henderson, which includes all wildpopulations with small fruits (0.5 to 10 g).

Clement et al.366

All cultivated populations are included in var. gasipaes and many wereclassified as landraces in three categories (Mora Urpí and Clement, 1988):microcarpa (fruits 10 to 20 g), mesocarpa (20 to 70 g) and macrocarpa (larger than70 g). At the time of the European conquest, this variety of landraces was widelydistributed throughout the humid Neotropics (Figure 18.3). Var. gasipaes exhibitshigher diversity in Amazonia (Henderson, 2000).

Within var. chichagui, Henderson suggested that there were three wildtypes: type 1 – very small fruits (0.9 to 1.6 cm long x 0.5 to 1.5 cm in diameter,weighing 0.5 to 1.5 g); type 2 – very small fruits (1.0 to 1.5 cm long x 1.0 to 1.4 cmin diameter, weighing 0.5 to 1.5 g); and type 3 – small fruits (1.5 to 2.9 cm long x1.4 to 2.8 cm in diameter, weighing 1.5 to 10 g). The considerable variability infruit size and disjunct distribution along the Andes (Figure 18.3) suggests thattype 3 could have been involved in the origin of the cultivated peach palm.

Figure 18.3. Distribution of Bactris gasipaes in the Neotropics at the time of theEuropean conquest, with the three types of var. chichagui (A;Clement et al., 2009) and the 14 primitive landraces of var. gasipaes(B; Cristo-Araújo, 2008): microcarpa –6. Tembe, 7. Juruá, 8. Pará;mesocarpa – 1. Rama, 2. Guatuso, 3. Utilis, 4. Tuira, 5. Cauca, 9.Solimões, 10. Pampa Hermosa, 11. Tigre, 12. Pastaza, 13. Inirida;macrocarpa – 14. Putumayo, 15. Vaupés.

The peach palm is diploid, with 30 chromosomes (Röser, 1999), althoughplants with 28 have also been reported in the literature. There are no reports onthe quantity of DNA in the nucleus. An analysis covering all the primitivelandraces and types of var. chichagui would be a worthwhile project, sinceerrors can occur in the karyotyping of domesticated plants.


Reproductive Biology

When well nourished, the peach palm can begin flowering in the fieldwithin three years, although it more commonly takes five years. In CentralAmazonia, the main flowering period starts in September, in the middle of the dryseason, with fruits ripening from the end of December to the beginning of March,in the middle of the rainy season. In years of high rainfall and when the mainharvest is not very abundant, the peach palm may flower a second time, producinga second small harvest in June and July, at the beginning of the dry season.Changes in the timing of the rainy and dry seasons also result in changes ofphenology, but no accurate analysis of these variations has been carried out.Note that the peach palm is seasonal, which is a serious limitation for industrialfruit processing.

The peach palm is pollinated mainly by weevils (Curculionidae) of thegenera Andranthobius (Central America) and Phyllotrox (Amazonia), and bysap-feeding beetles (Nitidulidae) of the genus Epurea. These small beetles havelimited flight endurance of between 100 and 200 m, rarely reaching 500 m. Largebeetles (Scarabaeidae) of the genus Cyclocephala have been reported in CostaRica, but so far have not be observed pollinating peach palm in Amazonia. If theflowers are not pollinated by insects, the wind and gravity can act as pollinatingagents, although they are less efficient. Efficient pollination is a problem inregions where the peach palm has been recently introduced, such as the AtlanticRainforest.

Although the peach palm is predominantly allogamous, self-pollinationcan occur between stems in the clump, which means that it could be more or lessfrequent. In addition, the traditional agricultural practice of planting a number ofopen-pollination seedlings from the same selected parent (half-sib progenies) inthe same field or home garden allows crossing between parents, resulting in bi-parental endogamy. The significant levels of endogamy in the peach palmpopulations of peasant and indigenous communities in Peru seem to be mainlyof this type (Cole et al., 2007).

Recently, Rodrigues (2007) investigated the reproductive system usingeight microsatellite loci in open pollinated progenies from three populations ofthe Pampa Hermosa landrace used in progeny trials. Estimates for crossing rateswere high for all peach palm populations, with 9 to 20 sources of pollen perprogeny, suggesting a mixed reproduction system with cross-fertilizationpredominant, probably due to the phenological stage (harvest peak) and floweringsynchronism during this stage, both aspects related to pollinator behavior.Significant endogamy was detected in all three populations, both uni-parentaland bi-parental. However, it is important to replicate these analyses at the

Clement et al.368

beginning and end of flowering to elucidate the variation in crossing rates andendogamy in this species, since variations in phenological synchrony can reducethe number of pollen donors and possibly increase endogamy rates.

Origin

The origin of the cultivated peach palm has been the subject ofspeculation for more than a century, since Richard Spruce (1871) suggested thatit could have originated in the Eastern Andes of Colombia. With the revision byHenderson (2000), it is obvious that domestication must have occurred withinthe areas of distribution of var. chichagui. However, type 2 exhibits a germinalpore pattern on the seed that is different from the other types of var. chichaguiand var. gasipaes, which would suggest that type 2 was not involved inoriginating the cultivated populations (Ferreira, 1999). Molecular analysissupports this conclusion (Hernández-Ugalde et al., 2008).

There are three hypotheses currently under discussion as to the originof this crop: the Colombian Andes, south-western Amazonia and multiple origins.In addition to Spruce’s speculations, the existence of numerous type 3populations of var. chichaqui in the Colombian Andes backs up the firsthypothesis. Carbonized seeds have been found at two archaeological sites inCosta Rica, dated at 2250-1650 years before present (BP), and at a site on thePacific coast of Colombia, dated at the same period (2190±60 years BP), whereasthe oldest evidence from Colombian Amazonia dates from 1080±40 years BP(Morcote-Rios and Bernal, 2001). However, there are only four sites with peachpalm remains and they are all very late. The peach palm exhibits sufficientdomestication-induced modification to justify dating its origin to the beginningof the Holocene (Clement, 1988) and not the late Holocene, as these sites suggest.

The possible origin of the peach palm in south-western Amazonia has beenthe subject of ongoing speculation since the beginning of the 20

th century. In this

region, there are numerous populations of the two types of var. chichagui which aremore likely to have been involved in originating the cultivated peach palm. All themolecular analyses conducted over the last decade have detected a clear-cutdifference between the populations cultivated in central and eastern Amazonia,including the populations all along the Madeira River, and those of western Amazonia,north and north-western South America and Central America (Rojas-Vargas et al.,1999; Rodrigues et al., 2004; Silva, 2004; Cristo-Araújo, 2008; Hernández-Ugalde etal., 2008). The most parsimonious interpretation of this division is a single origin inthe south-west, with dispersals to the north-east, along the Madeira and Amazonrivers, and to the north-west, as far as western Amazonia, northern South America


and southern Central America (Rodrigues et al., 2004). During this dispersal, thelikelihood of introgression with local type 3 wild populations is high, which wouldhelp explain the variability observed.

The considerable phenotypic variability of the peach palm can also beexplained by multiple origins (Hernández-Ugalde et al., 2008). Based on ananalysis covering numerous wild and cultivated populations and using fourmicrosatellite markers, Hernández and colleagues detected four distinct groups:i) three populations of type 2 var. chichagui from northern Colombia, unrelatedto the cultivated peach palm; ii) the cultivated populations along the Madeiraand Amazon rivers and a population of type 1 var. chichagui from the upperPurus river; iii) the cultivated and wild (type 3 var. chichagui) populations ofCentral America and the Cauca river valley in western Colombia; and iv) thecultivated and wild (types 1 and 3 var. chichagui) populations of westernAmazonia and northern South America to the east of the Andes. Based ongroups 2, 3 and 4, these authors suggest that there were at least threedomestication events. However, their argument is weakened by the omission ofthe cultivated population of Yurimaguas, in Peru, since when included in one ofthe groups, it combines with the populations of group 3 (Hernández-Ugalde,2005), creating a situation similar to that found by Rodrigues et al. (2004), inwhich the landraces of western Amazonia and Central America form a continuum.But this hypothesis cannot be discarded because there are other examples ofcrops originating from two or more domestication events in the Americas.

Domestication

The process of domesticating populations of plants has been describedby Clement et al. (Chapter 1). It is important to note that the response to selectiondepends on the selection intensity and the genetic isolation of new populationsoriginated by human propagation, as occurs in genetic improvement (Simmondsand Smartt, 1999). Selection intensity relates to the frequency of desirablephenotypes and isolation is related to dispersal beyond the natural distributionof species or to cultivation beyond the reach of natural pollinators. The degreeto which fruits are modified, as observed between the type 1 var. chichagui andthe Putumayo primitive landrace of var. gasipaes, is on the order of 2,000%,which suggests that domestication must have begun in the Holocene (Clement,1988). There are three hypotheses as to why the peach palm was domesticated,mentioned in the introduction to this chapter: for its wood, for its oily fruits andfor its starchy fruits. It is highly likely that all three are valid, but representdifferent stages during the domestication process (Clement et al., 2009).

Clement et al.370

The wood of the peach palm stem is preferred for making hunting andfishing tools, and for construction of artifacts, even house walls and flooring, andthese uses may be the reason why the peach palm was domesticated (Patiño,1989). The populations of var. chichagui are small and located near watercourses.Hunter-gatherer camps were also located near watercourses, where humansprobably discovered uses for the wood. It is unlikely that these humans propagatedthe peach palm, but cutting down a peach palm entails opening a small clearing towork in, which would increase the light available to the clump. This humanintervention could have had a positive effect on the reproductive success of theclumps, maintaining the abundance of peach palms, even though some stemswere cut down. Although the small fruits may not have attracted attention, whichis unlikely, the human groups could have taken peach palm seeds to other campswhere it did not grow naturally, thereby isolating new populations. The selectionintensity was probably zero, since any seed would produce a new clump withuseful wood, and isolation of populations probably occurred over many humangenerations. Changes in the phenotypes of the peach palm were unlikely, althoughnew populations probably exhibited low genetic variability due to the foundereffect and genetic drift in small, isolated populations.

The oily fruits would have been an important source of energy in a dietrequiring a high energy component. Over the last few years, Ana Vilacy Galúcio(Emílio Goeldi Museum, Belém, Pará, personal communication, 2006) and JorgeVivan (Federal University of Santa Catarina, personal communication, 2005)observed the preparation of wild peach palm in indigenous communities, Galúcioin Rondonia and Vivan in Acre. In Rondônia, type 1 fruits are cooked and eatenin the same way as cultivated peach palm fruit. In contrast, type 1 peach palmhas only 10-20% edible flesh in a fruit that weighs 1 g, as against 80-95% indomesticated fruits that weigh more than 20 g. In Acre, type 1 fruits are left tosoak in warm water, and then the flesh is mashed and separated from the seedand the juice squeezed and drunk. This form of preparation is very similar to themethod used for açaí (Euterpe oleracea, E. precatoria) and other Amazonianpalm fruits. The hunter-gatherers would also have prepared the peach palm inthis way, which changes the focal point of domestication. If humans took seedsfrom plants that produced an especially tasty juice to other camps, even withinthe natural distribution of peach palm, this seed migration would have allowedhybridization among separate populations, increasing the probability of findingeven more interesting variations. The most visible variations are the color of theexocarp and the size of the fruit. In wild peach palms, the red color is the norm,and yellow fruits are very rare, whereas in cultivated palms, yellow fruit is verycommon, reaching 20-30% in many populations. Of these two features, the mostimportant for selection is fruit size, since it opens new perspectives for


domesticating the plant. The easiest way of inducing a plant to produce largerfruits is through the accumulation of starch, rather than oil, which is much richerin energy. In other words, once the plant begins to be selected for its fruit, wecan say that this directly influenced the increase of starch in the mesocarp.

As it began to be used to make juice for regular consumption (alwaysafter cooking), the fruit increased in size, leading to an increase in starch content.When starch became more abundant than oil, it was possible to ferment the juiceinstead of letting it go rancid, and therefore new uses became possible. Innorthwestern South America and southern Central America, where the peachpalm was more important at the time of the European conquest, many ethnicgroups celebrated the peach palm harvest by feasting on cooked fruit and drinkingcaissuma (fermented peach palm juice). Nine months after these celebrations,the birth rate peaked (Patiño, 2002). Every year the peach palm harvest wasanticipated as a period of plenty, when people put on weight and became healthier(Patiño, 2002), since the fruit is rich in energy and beta-carotene, with somegood-quality protein (Yuyama et al., 1999, 2003). The degree to which the fruitwas modified in this region reached 2,000%, as mentioned earlier, mainly in thePutumayo and Vaupés landraces. These landraces can be seen as the apogee ofindigenous efforts to domesticate the peach palm, but not the only one.

A little to the south of the area in which the Putumayo landrace isfound, is the region of Yurimaguas, Peru, where the indigenous peoples alsoselected spineless plants, both spineless stems and spineless leaf petioles. Thisregion is home to the Pampa Hermosa landrace, the most important for peachpalm heart-of-palm agribusiness. Chávez et al. (1990) and Clement and Manshardt(2000) showed that the spineless traits of the peach palm are multigenic, withquantitative inheritance and low to medium heritability, which suggestsconsiderable and ongoing selection efforts to reach the high level of spinelessplants observed (60 to 80% in different populations around the Yurimaguas).Given this selection effort, which must have demanded high selection intensity,it is curious to observe that the Pampa Hermosa landrace is the most variable, inboth morphological and genetic terms (Mora Urpí and Clement, 1988; Clement etal., 1997; Rodrigues et al., 2004; Hernández-Ugalde et al., 2008). It is likely thatthis high level of variability is due to type 1 and 3 populations of var. chichaguiin the region that cross with the cultivated peach palm.

Landraces

A landrace is a set of domesticated, cultivated populations (meta-population), defined by a specific combination of original genetic resources and

Clement et al.372

associated ethnic history, with a defined geographic distribution, in whichsubpopulations are subject to continuous gene flow and genetic drift, and whosephenotypic traits are maintained by selection. In this case, primitive means“original” and not “backward”, as the word is sometimes used in the vernacular.There are a number of names for this concept of landrace, including local orCreole variety. In the case of peach palm, researchers differentiate landraces,since a single race occurs at any given location, but farmers do not make thisdistinction (which does not mean that the farmer is not aware of the variability ofthe peach palm in the region). This contrasts with the case of maize, for instance,in which the farmer uses and maintains different primitive landraces on hissmallholding, each with its local name (Louette, 2000).

At the beginning of the 1980s, a series of peach palm prospectingexpeditions were made in international Amazonia and the information obtainedwas combined with information from previous expeditions in Central America tobegin identifying, describing, classifying and mapping peach palm landraces(Mora Urpí and Clement, 1988). The peach palms on the Pacific coast of Colombiaand southern Central America are vegetatively more robust, with heavier andmore numerous leaves, and more spines on the stems than the peach palms inAmazonia and northern South America, which suggests a basic division in theracial hierarchy, with the Andes as the dividing line. On the next hierarchicallevel, fruit size is the differentiating factor, with races designated microcarpa(fruits from 10 to 20 g), mesocarpa (20 to 70 g) and macrocarpa (heavier than 70g) along a continuum of increasing weight. Fruit size is believed to be a reflectionof the degree of domestication, and therefore of the importance of the peachpalm to the humans associated with each landrace, especially in pre-Colombiantimes. Other fruit characteristics are also more varied between landraces thanwithin them, even where the phenotypic variation of the peach palm is verywide. A number of chemical and organoleptic characteristics of the fruit areimportant for human consumption and for this reason different races can beused as genetic bases for different objectives in breeding programs.

The initial definitions of the landraces were based on morphometriccharacterization. With the development of molecular genetics, it became possibleto perform genetic-molecular validation of the landraces, which also meant thatphylogenetic relations could be examined. Initially, RAPD markers were used toevaluate the hypothesis that there were Putumayo, Solimões and Pará landracesdistributed along the Solimões and Amazon rivers. The conclusions validatedthe genetic existence of the Putumayo and Pará landraces, with the Solimõeslandrace closely related to the Putumayo race (Sousa et al., 2001). Next, Rodrigueset al. (2004) evaluated seven more landraces, four in Amazonia (Pará, Putumayo,Solimões, Pampa Hermosa) and three in Central America (Utilis, Tuíra, Guatuso).


Molecular analysis validated the Pará, Pampa Hermosa and Putumayo landracesin Amazonia, with the Solimões populations belonging to the Putumayo landrace.In Central America, only the Utilis landrace was validated, with the Tuíra andGuatuso populations belonging to Utilis. However, a recent morphometricanalysis suggests that the Solimões race should be kept separate but morelimited in size than originally proposed (Martel et al., 2003). Silva (2004) confirmedthe validations of Rodrigues et al. (2004) and showed the validity of the Juruálandrace and its relationships with the other three western races (Pampa Hermosa,Putumayo and Utilis). The other landraces evaluated by Silva (Vaupés, Caucaand Inirida) were represented by only small numbers of individuals, barring amore precise conclusion, but always showing relationships with the westernlandraces. Recently, Cristo-Araújo (2008; Cristo-Araújo et al., 2010) finalized ananalysis of the landraces present in the INPA Peach Palm Active GermplasmBank (BAG-Pupunha) in Manaus, confirming the results previously obtainedand revealing interesting relationships between populations in northern Boliviaand the Pará landrace, and those in southeastern Peru and the western Amazonianlandraces. Four hybrid populations (Belém, Manaus, Iquitos and Yurimaguas)were also characterized using RAPD, and it was observed that these populationsdid not exhibit greater genetic variability than the landraces, although those inManaus exhibited greater heterozygosity than the others, probably because it isa meeting point for eastern and western dispersals (Santos et al., in press).

In view of the importance of the Pampa Hermosa landrace for theproduction of hearts-of-palm, these analyses were extended with the aim ofverifying the existence of other landraces around the Pampa Hermosa landrace,as suggested by Mora-Urpí and Clement (1988), based on the morphometricvariability observed in the Yurimaguas region. Silva et al. (2003) evaluated threepopulations around the Yurimaguas in Peru and verified that they did not differfrom the Pampa Hermosa landrace. Using AFLP, Adin et al. (2004) comparedgenetic diversity among the populations along the Paranapura and Cuiparillorivers and detected little divergence and high gene flow, providing support forthe hypothesis that they are populations of the Pampa Hermosa landrace, whichwas confirmed by Rodrigues (2007) when she evaluated the diversity and geneticstructure of the Pampa Hermosa landrace using eight microsatellite loci.

Genetic Resource Collections

Numerous collections of genetic resources have been set up to begingenetic improvement of the peach palm. Unfortunately, it has proved mucheasier to set up a collection than to maintain, characterize, evaluate and use it.

Clement et al.374

The story behind the work on peach palm collections is as varied as the peachpalm itself, with some old collections still active, other new and active collections,and others completely extinct, but even the active collections have contributedlittle to the development of the crop (Clement et al., 2004). The first two peachpalm collections in Brazil were created at the Agronomic Institute of the North(IAN) in Belém, now extinct, and at the Agronomic Institute of Campinas (IAC).

Work to set up the Active Germplasm Bank at the IAC began in 1973, withthe introduction of peach palm accessions from Brazilian Amazonia, Costa Ricaand Peru. New accessions were acquired in the 1980s, mainly of spineless peachpalm from the Yurimaguas region of Peru, sent by INPA. After plant selection inthe greenhouse based on the absence of spines on the petiole/rachis, the numberof live leaves and plantlet diameter just above the meristem, a seed productionsite was set up in Ubatuba, state of São Paulo, at a spacing of 5 x 4 m. Althoughits genetic base was restricted, this site increased the number of peach palmsfrom the Yurimaguas population. The seeds were distributed in small quantitiesto some farmers, soon providing evidence of the superiority of the Peruvianmaterial for producing hearts-of-palm and its advantages in comparison to thespecies traditionally used for palm heart production (Euterpe edulis and E.oleracea) in the state of São Paulo. The third and last prospecting expedition tothe Yurimaguas was made in 1990, with the introduction of seeds of 332 progenies,expanding the genetic base available. Although this collection concentrates onspecimens from the same population, the way it was established and evaluatedhas made it the main working collection at the IAC.

The Active Germplasm Bank (AGB) at INPA (National Research Institutefor Amazonia) was set up in 1976, expanded considerably during the 1980s toreach 455 accessions, then stagnated at the beginning of the 1990s. Work onreviving it began in 1995 and is still under way. The AGB accumulated accessionsfrom collections made during numerous peach palm prospecting expeditions,but its representativeness is flawed insofar as it contains mainly Brazilianlandraces and hybrid populations, and the Pampa Hermosa landrace. It wasconceived as a collection for both conservation (at a time when the nativegenetic resources were thought to be under threat of genetic erosion) andbreeding. However, its design does not allow experimental work for comparingaccessions because accessions of interest are surrounded by accessions oflittle interest, making mass selection difficult and requiring controlled pollination.As a result, the AGB has not contributed to breeding programs (Clement et al.,2004; Clement et al., 2005; van Leeuwen et al., 2005), although it has beenfundamental in understanding the domestication of the peach palm.

Since this is a living collection, AGB maintenance costs are high andfunding to characterize and evaluate accessions is increasingly scarce, limiting


its use. One way of improving conservation, concentrating efforts and cross-referencing information to promote the efficient use of the AGB is to create aCore Collection (CC), consisting of a set of accessions representing at least 70%of the genetic diversity of the whole collection, with minimum duplication(Cordeiro and Abadie, 2007; Johnson and Hodgkin, 1990). To design a CC,accessions were stratified into two groups based on the level of domestication(wild and cultivated), then the cultivated accessions were stratified into landracesand hybrid populations. Landraces and populations that are poorly representedin the AGB were allocated to the CC in proportion to their number: Juruá (2),Cauca (2), Guatuso (2), Pastaza (1), Tuira (1), Utilis (2), Vaupés (2), undesignatedpopulations (5), and wild populations types 1 and 3. Well-represented landracesand populations were allocated based on the logarithm of their number: Pará (5),Putumayo (4), Solimões (3), Pampa Hermosa (4) and hybrid populations (3).Accessions within landraces and populations were selected based on thedivergence observed in published Jaccard or Dice similarity matrices. Therecently-created CC was put together with 10% of the entire collection, with 28landrace accessions, three hybrid populations, five undesignated populationsand four wild populations. The majority of accessions consist of nine plants.The CC is thought to contain at least 80% of the AGB’s variability (Cristo-Araújo, 2008). This core collection will facilitate further efforts to characterizeand evaluate accessions, and to adequately plan programs aimed at conservationor breeding, making the collection more useful.

The research usually carried out on collections, such as morphometricand DNA studies, can also be conducted using comparative trials. At present,there are four good progeny trials under way, each associated with one of thebreeding projects discussed below: Embrapa Amazônia Oriental’s fruit productiontrials, IAC’s palm heart trials, INPA’s palm heart trials and palm heart trials organizedby the Embrapa breeding network, headed by Embrapa Florestas.

Breeding for Fruit – National Research Institutefor Amazonia (INPA)

Since the Peach Palm AGB was set up, INPA researchers have distributedseeds to producers in different parts of Amazonia and Brazil. However, the resultshave not lived up to expectations because the recipients of the seeds wereunder the impression that they were improved material, whereas in reality theyreceived heterogeneous material obtained from crossings at the AGB. At theend of the 1970s, INPA researchers planned breeding work on the peach palmwith four different fruit-related objectives: the whole fruit for human consumption,

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coarse flour for animal feed, flour and starch for baking, and extraction of oil. Thebreeding strategy adopted by INPA researchers allowed for the setting up andmaintenance of an AGB representing species variability, the characterizationand evaluation of this bank (more than 3,500 palms at its peak), controlled crossingsof the best stock and progeny trials at the experimental station. This strategywould have taken a long time and required ongoing funding, and therefore wasnot implemented. In the mid-1980s, INPA received funding from FINEP (Studyand Project Finance Agency) to begin a breeding project with two objectives:hearts-of-palm, and starchy fruit for human consumption and industrialprocessing. To achieve the first objective, efforts centered on the Pampa Hermosalandrace, with a few Putumayo plants. For the second objective, the researchersworked mainly with the Putumayo landrace, since this is the starchiest landrace,with good representation in the AGB and in other INPA plantations. However, atthe end of the 1980s, economic problems in Brazil led to the abandonment of theproject. Looking back on the objectives of this project, today it is clear that thefirst was well-suited to the situation, since the demand for peach palm seeds forproducing hearts-of-palm was already growing in Brazil. However, the secondwas not, because the market assessment was incorrect and not based onconsumer survey data. At present, breeding programs are aimed only at producingfruit for human consumption, the only commercial peach palm activity.

At the beginning of the millennium, a survey was conducted on consumerdemand in Manaus (Clement e Santos, 2002). Of those interviewed in the Manauscentral market, 40% did not like peach palm, including the Amazonas nativepopulation and people who had been living there for some time. Among theconsumers who liked peach palm, few consumed more than a bunch per weekduring the three-month harvest period, around half of them preferred the oily fruitand 40% liked a well-balanced oily-starchy fruit. The landraces that satisfy thefirst group are Pará and Juruá, and for the second group, the Solimões is better,especially the fruit from the eastern region which is slightly oilier. Note that thePutumayo landrace, previously used by the INPA, did not have any enthusiasts inManaus. INPA researchers are planning to use a participatory breeding strategy(see below). Against this backdrop, Embrapa Amazônia Oriental began new workaimed at satisfying demand in Belém, which is similar to that in Manaus.

Breeding for Fruit – Embrapa AmazôniaOriental

One of the main obstacles to boosting peach palm consumption in thestate of Pará is the shortage of high-quality fruits with standard traits, such as


oil content, quantity of fibers, pulp yield and fruit moisture content. The consumerdoes not have access to fruits with the desired characteristics simply becausethe producer generally uses unselected seeds for planting, and becauseresearchers have not yet developed desirable cultivars. Producers thereforeplace fruit of dubious quality for the market.

To supply the market with a product that satisfies the requirements of theproducer, wholesaler and consumer, Embrapa Amazônia Oriental began a peachpalm genetic improvement program to produce quality fruits. In April and Mayof 2005, 80 promising bunches of peach palm fruit were acquired from thenortheast of Para and along the Madeira river, in Amazonas, based on thefollowing bunch selection criteria: medium-size to large fruits (for the Parálandrace, weight ranges from 20 to 40 g), red in color (indicating higher beta-carotene content than yellow or green fruit), with low moisture content in thepulp and good pulp yield.

After removing the seeds, the fruits were cooked, subjected to taste andacceptability testing, and assessed by scores from 5 to 9 for oil content, pulpfiber and flavor. A total of 50 open-pollinated progenies obtained a score higherthan eight and their seeds were germinated to provide seedlings and set up atrial, which began in February 2006. In the trial, the seedlings were laid out in 15fully randomized blocks with one plant per plot in each block (single-tree plot).The progenies are randomized within the blocks to prevent endogamic depressiondue to selection as the seed orchard is formed.

Participatory Breeding for Fruit

The fruits offered for sale in Manaus are extremely diverse in terms ofcolor, size, shape and quality, with a high percentage of poor fruits. Since thereare no commercial varieties, the buyer does not know how to choose the rightfruit. He will only know whether he has made a good buy when he arrives homeand tries the cooked fruit. The result is often disappointing. As a consequence,peach palm fruit is not seeing the increases in sales that other regional fruits areexperiencing, stemming from the considerable increase in the urban populationin Amazonia.

The lack of results from the INPA breeding program led to a review ofstrategy. Since there were no improved varieties, it was essential to produce oneas quickly as possible. A strategy based on recurrent selection could providethe first improved seeds within six to eight years. A simple and fast programbegins by selecting parents with superior traits in producer areas, and then

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setting up trials and comparing the progenies of these parents. These trials canalso be used for producing improved seeds. It is not necessary to characterizethe trees based on numerous descriptors. Yield and quality parameters aresufficient (van Leeuwen, 2006).

Maintaining long-term trials at experimental stations can be fraught withproblems, since research projects rarely extend beyond a few years. As theproject is coming to completion, funding for maintaining the trials can run out.For this reason, it is often better to place some or all progeny trials in the handsof farmers, who will set them up and maintain them. There are other advantagesin doing this, including the help of the farmer in evaluating the material andhaving data on actual production conditions. A program of this kind has beenimplemented in Peru since 1996 (Cornelius et al., 2006). Further details of a proposalbased on recurrent selection, together with issues such as trial locations inproducer areas, trial design, farmer collaboration and the necessary marketingand commercialization strategy for the new variety of fruit, are discussed by vanLeeuwen (2006).

Breeding for Hearts-of-Palm – CampinasAgronomic Institute (IAC)

Peach palm seeds were introduced by the IAC around 1940. Seedlingsfrom these initial seeds were planted in the fields of a few established ExperimentalStations. However, research on the peach palm began only in 1973, based on anencouraging paper by Camacho and Soria (1970) that showed the potential ofthe peach palm for producing hearts-of-palm. Using seeds imported from CostaRica, Peru and Brazilian Amazonia, experimental plots were set up at five locationscovering the diverse edaphic and climatic conditions prevalent in the state ofSão Paulo. Many of these plots are still flourishing, and among these some 30%no longer have spines on the stem and petiole/rachis, and produce fruit inabundance.

At the beginning of the 1980s, new peach palm stock was introducedinto Ubatuba. The Paraíba Valley region in which this collection is located waschosen because of its ideal conditions for producing hearts-of-palm. Over theyears, however, the region has shown little promise for producing seeds. This iswhy the studies conducted have focused on flowering and the entomofaunathat visit the inflorescence, but have yet to produce results.

It is essential to point out that, from this decade onwards, the majority ofstudies carried out at the IAC, not just on breeding but also for upgrading the


peach palm crop to produce hearts-of-palm in the state of São Paulo, wereconducted and coordinated by Dr. Marilene Leão Alves Bovi. These studies,combined with the easy adaptation and early production of hearts-of-palm, aswell as increased pressure and environmental awareness- in relation to PalmiteiroJuçara (Euterpe edulis Mart.) extractive activities, resulted in the peach palmbecoming the most cultivated species for heart-of-palm production. Bycomparison with 1990, 2005 saw a drop of 71% in hearts-of-palm extracted fromnative palms, and an increase in the heart-of-palm growing area by a factor ofalmost 90 (Anefalos et al., 2007). Also worthy of note is the fact that this cultivatedarea is almost entirely planted with peach palm.

The third and last prospecting expedition for material from the Yurimaguaspopulation took place in 1990, thanks to a joint project involving INPA, IAC andSão Paulo State University (UNESP). The material collected was used to set upfour germplasm banks consisting of half-sib progenies in two separate regions:the coastal region and the Paulista Plateau (Planalto Paulista). As a breedingstrategy, in parallel with the germplasm banks, progeny trials were set up indifferent farming regions in the state, totaling seven trials on the north coast(307 progenies), three on the south coast (103 progenies) and an experiment onthe Paulista Plateau (42 progenies). In this way, at the same time as the accessionsare maintained and evaluated in the germplasm banks, the progeny trials can beused to generate estimates of genetic parameters to help define selectionstrategies and to assess the expected gain from each selection cycle for eachtrait studied. These trials were set up using commercial spacing of 2 x 1 m, in fullyrandomized blocks, with between 9 and 12 plants per plot and without offshootmanagement. The material was evaluated in the greenhouse and later in the field,taking account of traits related to growth rate, the number of offshoots, theabsence of spines on the petiole/rachis and stem, as well as a few componentsdirectly related to heart-of-palm production. Based on these studies, Bovi andcolleagues evaluated traits for indirect selection of peach palms and verified thepossibility of early selection. Taking the traits related to growth, production ofoffshoots and hearts-of-palm, the best progenies were selected and are beingcultivated to produce seeds. The quality of the hearts-of-palm from some ofthese progenies was also evaluated in collaboration with the Food TechnologyInstitute (ITA).

With regard to the germplasm banks, it will only be possible to use theirgenetic variability if the material is properly characterized and evaluated. Forthis purpose, the working collection is concentrated at the bank set up inPindorama (21º 13’S and 48º 56’O, 560 m elevation), São Paulo state, whichcontains 332 progenies (1991 plants) from the collections in Yurimaguas, Peru,including 1,048 collected from the Huallaga river region, 444 from the Cuiparillo

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river, 343 from the Paranapura river and 156 from the Shanusi river. The evaluationswere focused on morpho-agronomic descriptors of interest for cropping: stemheight and diameter, number of offshoots, number and dimensions of leaves,shape and size of floral spathes, number, dimensions and color of male andfemale flowers, number, dimensions and color of fruits, and number, size andshape of seeds. Of the plants in the bank, only 337 are in the reproductive phaseand could be characterized. The process of characterization will proceed as theplants produce fruits.

Characterizing the germplasm bank and the use of uni- and multi-variateanalysis opened up a range of possibilities (Gomes, 2007), including a) theformation of accession groups based on genetic dissimilarity to define interestingcrossings, b) selection of plant groups with favorable traits for improving thespecies, in particular plants from Huallaga, Cuiparillo and Paranapura, and c)quantification of the percentage variability for each descriptor so that descriptorswith little relevance could be discarded (49% in the juvenile phase, 22% in thereproductive phase), cutting the workload, especially in terms of additionalreproductive phase evaluations.

The IAC is still conducting participatory breeding with collaboratingcompanies, using stratified mass selection in commercial palm heart crops. Thisinvolves using a selection index of no more than 20% and the criteria outlinedabove. Plants selected as stock have been evaluated periodically to verifysuperiority, based on the number of fertile fruits per cluster, germination speedindex and percentage, and the vigor of the plants obtained.

Breeding for Hearts-of-Palm – INPA

The objective of peach palm breeding for hearts-of-palm at the INPA is toobtain progenies with rapid growth, more than four rapidly growing offshootsper year, high frequency of hearts-of-palm longer than 45 cm and plants withoutspines on the stems and leaf petioles/rachis. This ideotype is narrower than theone described by Mora Urpí et al. (1997), which should speed up the geneticselection process. The program began in 1991 with 391 spineless accessionscollected in the Yurimaguas region in Peru along four river systems: Cuiparillo,Huallaga, Paranapura and Shanusi (Clement et al., 2001). There are minormorphometric (Yuyama et al., 2002) and genetic (G

ST = 0.019; Rodrigues, 2007)

differences between these populations, which means that they all belong to thePampa Hermosa landrace. In 1993, 257 accessions were planted at the INPATropical Fruit Experiment Station in fully randomized blocks with three replicationsof 20 plants per plot spaced at 2 x 1 m. Agronomic characterization and evaluation


began in 1994 and was completed in 2001. It was based on the descriptors ofClement and Bovi (2000), with a few additions. This information provided a basisfor inter- and intra-progeny selection, using the following characteristics ascriteria: i) inter-progeny selection – precocity, offshoot production (more thanfour offshoots/plant), heart-of-palm length (higher frequency of hearts over 45,54 and 63 cm long); ii) intra-progeny selection – repeatability of heart-of-palmlength and weight (45 cm and 200 g), with a minimum length of 45 cm, and theabsence of spines on the petiole, rachis and stem, with a minimum of four perfectoffshoots/plant. In the initial selection, 863 plants were identified with longhearts (Table 18.2) and subsequently short-listed to 132 plants based on otherinformation, since many of them did not maintain offshoot production, died forother reasons or exhibited spines later on.

Next, the majority of plants selected were genotyped using eightmicrosatellite markers, together with a representative sample of rejected plantsfrom the same progenies (Rodrigues, 2007). The progenies exhibited high geneticvariability: 82.7% intra-progeny, 16% inter-progeny, and only 1.3% among thepopulations defined according to river systems. The dendrogram of sharedallele distances among the progenies shows high genetic affinity, with fewdivergent progenies. The formation of progeny groups is independent ofgeographic origin, although population groupings show greater affinity betweenthe Shanusi and Huallaga rivers, and between the Paranapura and Cuiparillo.This suggests that there is almost no population structure within the PampaHermosa landrace. This information will be useful for determining controlledcrossings of progenies within each population based on morphometriccharacteristics and genetic divergences, with the aim of maximizing intra-population heterozygosity, and then inter-population heterozygosity (recurrentselection) to favor the formation of heterotic groups that could achieve highergenetic gains.

Outstanding Progeny Traits

Generally speaking, the progenies of the Shanusi River populationexhibited greater vigor, apparently due to the greater number of green leavesand length of the rachis on the third leaf, and the precocity and greater diameterof the stem at the moment of heart-of-palm extraction. However, the sheath waslonger in the Paranapura River population and shorter in the Huallaga riverpopulation. Once the heart length is defined, it can be confirmed that the numberof green leaves is not directly correlated with palm heart length.

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The populations from the Cuiparillo, Huallaga and Paranapura Rivershad a higher progeny selection rate, and more plants were selected from withinthe progenies (Table 18.2). These progenies exhibited higher heterozygosityindices and lower average endogamy coefficients, verified by means of geneticanalysis. On the other hand, the Shanusi River population exhibited lowerpercentages of selected progenies and plants, and lower heterozygosity andhigher endogamy. Among the plants selected, those from the Paranapura Riverexhibited a higher percentage of palm hearts longer than 63 cm, whereas thosefrom the Shanusi River had a higher percentage of palm hearts between 54 and63 cm. It is still too early to confirm that these genetic parameters are decisive,since they are based on only eight microsatellite markers, but they do suggestthat this information could be of use to the program in the future.

Table 18.2. Progenies and plants initially selected over the period from 1994 to2001, based on heart-of-palm length in a trial of 257 progenies (n=60/progeny) of peach palm from the Pampa Hermosa landrace kept atthe National Research Institute for Amazonia (INPA) ExperimentalStation.

Cuiparrillo Huallaga Paranapura ShanusiRiver River River River

Total progenies 65 128 47 17

Selected progenies 49 (75%) 98 (77%) 36 (77%) 10 (59%)

Selected plants 230 (6%) 441 (6%) 158 (6%) 34 (3%)

> 45 and < 54 cm 175 (76%) 335 (76%) 91 (58%) 20 (59%)

> 54 and < 63 cm 53 (23%) 97 (22%) 61 (39%) 14 (41%)

> 63 cm 2 (1%) 9 (2%) 6 (4%) 0

Obs. heterozygosity 0.680 0.690 0.740 0.640

Endogamy coefficient 0.132 0.124 0.111 0.190

The observed heterozygosities and endogamy coefficients are representative samplesfrom the selected progenies, analyzed using eight microsatellite markers (Rodrigues,2007).

Future Strategies

The next stage in improving the peach palm will be hybridization betweenthe best selected progeny plants that are genetically divergent to determinewhether there is any hybrid vigor due to different combinations of rare alleles.


These hybridizations will also allow the evaluation of any effect of heterosis, aswell as general and specific combining abilities. The same plants will be self-pollinated and planted next to the progeny competition trials. In parallel withthis study, pest and disease resistance will also be identified.

If hybrid vigor is not observed in this trial, the spineless plants in thepeach palm AGB will be evaluated for use as sources of new alleles via controlledpollination. The introduction of new alleles presents some risks, since the plantscannot be evaluated for heart-of-palm length, although the length of internodesis associated with rapid growth and the number of offshoots is known. The mostdivergent landrace is Pará (Rodrigues et al., 2004), but it unfortunately exhibitsan extremely low proportion of spineless plants. Next comes the Utilis landrace,and in particular the Costa Rican Guatuso population.

Later, the project could include the other traits suggested by Mora Urpíet al. (1997): leaves with erect petioles and rachis, and also erect leaflets, high netassimilation rate, initial harvest at less than 12 months (based on a plant heightof 1.5 m), second harvest between six and eight months later, and hearts-of-palmwith a white to creamy in color, good flavored hearts that do not cause irritationwhen eaten fresh (absence of calcium oxalate crystals). The first trait shouldallow higher crop density and, when combined with the second, should makethe third and fourth relatively easy to achieve. The traits relating to appearanceand organoleptic quality are important for the fresh hearts-of-palm market, whichis growing rapidly in Brazil.

Breeding for Hearts-of-Palm – Embrapa

Genetic resources within the Embrapa Plant Network

Embrapa has a collection of peach palm germplasm from the populationsof Yurimaguas, Peru, consisting of alleles from the Pampa Hermosa, Putumayoand other unidentified landraces with 60 to 80% spineless plants, and fromBenjamin Constant, Amazonas, Brazil, of the Putumayo landrace with 15 to 25%spineless plants (Clement, 1988). However, the germplasm from BenjaminConstant at Embrapa has been improved for absence of spines and vigor overtwo generations, the first at the INPA and the second within the RECA Project,a producer settlement in Extrema, Rondônia. The second improved generation atBenjamin Constant was the outcome of a partnership between Embrapa Acreand the RECA Project. The stock selected is in producer areas within the RECAProject and provides a source of seeds for commercial planting in Brazil. Thisstock produces on average 7.4% individuals with spines on the stem (Kalil Filho

Clement et al.384

et al., 2001). The Embrapa Centers also hold other peach palm genetic resources(Table 18.3).

Table 18.3. Open-pollinated progenies of spineless peach palm and the year inwhich they were introduced at the Embrapa Center involved in thenational progeny competition trials.

Embrapa Centers Yurimaguas Year RECA Project Year Others Year

Acre 20 2005 100 2002

Amapá 31¹, 64 1999 100 1999

Amazônia Ocidental 31¹ 82 2005

Amazônia Oriental 96 2005 95³ 1985

Florestas 40², 95 2005

Rondônia 50 2005

Roraima 102 20051 From the original introduction at INPA (1980) by Mr. Imar César de Araújo, who carriedout two generations of mass selection before handing over the progenies to EmbrapaAmazônia Ocidental, where mass selection was carried out to produce the third generation.The progenies were later sent to Embrapa Amapá, in 1997.

2 Introduced in 2001.3 Prospecting for pre-Colombian fruit trees, samples were taken from the municipalities ofFonte Boa, Tefé and Tonantins, in the state of Amazonas, collected for starchy fruit quality

and absence of spines.

Progeny trials

Various Embrapa Centers began breeding projects for hearts-of-palm atdifferent times. At Amapá, the first 31 Yurimaguas progenies exhibited low additivegenetic variance, with an estimated genetic gain of 8.4% in the short term (FariasNeto and Resende, 2001), which is not surprising given their selection history.The other 64 Yurimaguas and 100 Benjamin Constant progenies exhibited highgenetic variability for vigor traits, and the Benjamin Constant population exhibitedhigher genetic variability than the Yurimaguas population (Farias Neto andBianchetti, 2001). These progenies are still being evaluated. The repeatabilityestimates obtained show low uniformity in the superiority of progenies from oneevaluation to another, so that for the plant height trait, three evaluations arenecessary for prediction reliability of 80%. For traits related to diameter at breastheight and heart-of-palm weight, six evaluations are necessary for selectionwith the same level of reliability (Farias Neto et al., 2002).


After helping the RECA Project select parent plants, Embrapa Acre setup a trial with 100 progenies to evaluate their quality and select the best individualsfor heart-of-palm production. The Yurimaguas trial was then set up and both areunder evaluation.

In Paraná, the breeding program started in 2001, when a trial was set upwith 26 of the 40 progenies in Morretes (coastal region), 17 in Tagaçaba (coastalregion), 40 in Londrina (northeast) and 23 in Cidade Gaúcha (northwest). Theevaluations and analyses show that: i) average growth, offshoot productionand heart-of-palm production on the coast were higher than in Londrina, as weresurvival averages; ii) progeny-environment interactions (Londrina and Morretes)were complex, with low correlation between locations, indicating the need forlocal breeding; iii) genetic gains of around 30% were obtained for traits relatedto vigor, offshoot production and survival; iv) when subjected to a selectionintensity of 10%, the progenies exhibited genetic gains of 25% for height, 16.7 to24.5% for diameter, and 22.4 to 55.3% for number of offshoots; v) a substantialincrease in genetic gains, ranging from 69.9% to 121.8%, was obtained by indirectselection for heart-of-palm production and combining the criteria for height,diameter and number of offshoots (edible stem volume); and vi) the geneticcorrelation between height and diameter was 89.5% in Morretes and 87.85% inLondrina, between height and number of offshoots 31.85% in Morretes and65.56% in Londrina, and between diameter and number of offshoots 25.64% inMorretes and 75.61% in Londrina. Based on this information, the first seedproduction areas were set up, resulting in a selection process for 279 peachpalms in Londrina and 270 in Morretes.

The Peach Palm Breeding Network inAmazonia, Espírito Santo and Paraná

In 2004, a new project was initiated, the Macroprogram 2 Project, with theintroduction of new progenies from the Benjamin Constant population and afurther 20 progenies from Yurimaguas, Peru, as well as the evaluation of the progenytrials already set up in Amapá, Acre and Paraná. This new project therefore unifiedthe separate breeding programs under way at various Embrapa Centers into asingle program, culminating in a peach palm breeding network extending over thestates of Amazônia, Espírito Santo and Paraná, and involving 24 researchers. Thebreeding strategy for all Embrapa genetic improvement programs involves intra-population recurrent selection. The current phase of the networked program entailsevaluating and selecting superior genotypes for heart-of-palm production andsetting up improved peach palm seed production areas in each state, expected tobe completed within the next three years (by 2011).

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The Next Ten Years

Location- and species-adaptive genes for tolerance to environmentalstresses should be detected in the future by the Embrapa Breeding Network.These genes will be important for adapting the peach palm crop to the colder,drier, altered environment resulting from global warming. In addition, they willbe detected and concentrated in progenies with superior performance inproducing hearts-of-palm in different environments.

Crossings of superior genotypes from the improved Yurimaguas andBenjamin Constant populations, with wider genetic divergence, will lead to furthergains in heart-of-palm productivity for vigorous inter-population hybrids.Production of improved seeds from these populations will increase cropproductivity, and therefore economic viability, especially for small farmers (themost common) within the peach palm heart agribusiness, since it will increasefarmers’ incomes and help improve quality of life. Heart-of-palm productivityunder local conditions is expected to rise from 900 g tender palm heart (currentaverage weight for commercial crops) to 1,500 g per stem. Per hectare productionshould increase from 4.5 to 7.5 metric tons on average. Productivity could bedoubled if rooted offshoot clones could be produced and a commercial protocolestablished for tissue cultures.

The results to be obtained from the project developed by EmbrapaAmazônia Oriental should provide adapted, productive peach palm seedsimproved for fruit, capable of promoting greater crop uniformity and satisfyingthe requirements of producers, wholesalers and consumers. Selection ofindividuals for propagation from seed should be based on the genetic values ofselection candidates, which depend only on additive gene effects.

In allogamous species breeding programs, the search for inter- and intra-population genetic divergence could provide a basis for crossings to expandgenetic variability in segregating populations, creating heterotic groups. Sincethe progenies in the Yurimaguas trials exhibit a close genetic relationship,although with some of them more divergent than others, molecular marker-assistedselection could bring considerable benefits. Molecular analyses in the INPAtrial are being expanded, using more microsatellite markers to improve analysisof relations between heterozygosity, vigor and specific traits related to palmheart production. The inclusion of progenies without selected plants will alsoenhance our understanding of the genetic diversity of the Pampa Hermosalandrace in general.


Acknowledgements

This chapter is dedicated to our colleague Marilene Leão Alves Bovi(Agronomist, M.S., PhD., researcher at the Agronomic Institute’s HorticulturalCenter, Campinas, São Paulo, Brazil), for her significant contributions to researchon the peach palm over three decades, before her untimely death in 2006.

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Domestication and breeding of peach palm

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