Bactris Gasipaes

Pejibaye or peach palm (Bactris gasipaes, Kunth) was introduced into Hawai'i to supply the gourmet market with fresh heart of palm. New crop introduction requires evaluation of crop adaptation to its new environment and planning for future development, including genetic improvement. Leaf number of open pollinated Benjamin  Constant (Putumayo landrace) progenies was lower at harvest (6-8) than elsewhere (8-10), and offshoot number
dropped dramatically from first harvest (6.5) to second harvest (2). Allometric equations for estimating whole plant leaf area and biomass were developed, using height and leaf number predictors. No significant plant population (density) effects on individual plant dimensions or growth were found over the range of 3333 to 6666 plants/ha. Relative growth rate (RGR) and unit leaf rate (Ea) between nursery and first harvest were highly correlated (r = -0.99 and -0.95, respectively) with earliness (days to harvest). The early progenies partitioned photoassimilates differently: two had high Ea, while one had moderate Ea and partitioned preferentially to leaf area, resulting in a higher leaf area ratio. Heart of palm yields were close to 900 kg/ha after 12 months of harvest and 1400 kg/ha after 18 months, both comparable to tropical American yields. When edible stem and leaf were added to the yields, these increased to 2.8 and 4.5 t/ha of marketable product, respectively. Quantitative genetic analysis of growth parameters suggested high levels of inbreeding in the germplasm studied, since the narrow-sense heritabilities were double those observed in other perennials. Additive genetic variances for RGR and earliness suggested the potential for significant response to selection, but phenotypic variation varied depending on the interval over which RGR was estimated. The lowest estimate of RGR (over an entire development phase) provided the smallest response to selection but is similar to the response observed in other
crops. Allozyme heterozygosity was remarkably low, ranging from 0.038 to 0.099, with a mean of 0.074, on par with inbred crops, rather than outbreeders. There was a lack of correlation between allozyme heterozygosity and growth parameters and morphological traits.

The Peach palm Active Germplasm Bank has abundant genetic diversity in its holdings. Because it is a live collection, maintenance, characterization and evaluation are expensive, restricting its use. One way to promote more efficient use is to create a Core Collection, a set of accessions with at least 70% of the genetic diversity of the full collection with minimal repetition. The available geographic, molecular marker (RAPD) and morphometric information was systematized and the populations were stratified into wild and domesticated. The Core Collection consists of 10% of the entire collection: 31 accessions of landraces, 5 accessions of non-designated populations and 4 accessions of wild populations. The Core has 92% of the molecular polymorphism and 95% of the heterozygosity of the full collection, with minimal divergence between them by molecular variance. The Core is already receiving priority maintenance, which will contribute to long term conservation.

Peach palm is a native Amazonian fruit species with broad genetic diversity in its wild and domesticated populations. This study completed the molecular characterization and genetic analysis of the accessions in the Peach Palm genebank, Brazil, using RAPD markers. Eight primers generated 132 markers, of which 128 were polymorphic. Average heterozigosity was 0.35, with 95.5% polymorphism. The genetic variability within the landraces (H S) was 0.32, while genetic divergence (G ST) was 0.09 among the four well-represented landraces. The average gene flow was 5.0, with high gene flow between the Solimões and Pampa Hermosa (Nm=10.07), and between the Putumayo and Solimões (Nm=10.73) landraces, all western landraces. The dendrograms of the Nei's and Rogers' genetic distances of the well-represented landraces presented similar groupings to previous analyses, with the Solimões, Putumayo and Pampa Hermosa landraces grouped together in western Amazonia, and distant from the Pará landrace in central and eastern Amazonia.

Contents
1 Introduction 7
2 Taxonomy, nomenclature and geographical distribution 8
2.1 Taxonomy 8
2.2 Botanical and vernacular names 10
2.3 Geographical distribution 10
3 Description of the cultivated species 12
3.1 Morphological description 12
3.2 Ecology 14
3.3 Development of the stem and inflorescence 16
3.4 Reproductive biology 17
4 Uses and properties 20
4.1 Traditional food products 20
4.2 Current food products and their properties 20
4.3 Minor products 24
5 Origin and domestication 26
6 Genetic resources 28
6.1 Diversity among and within landraces 28
6.2 Existing germplasm banks 30
6.3 Useful and unusual phenotypes 32
6.4 Genetic erosion 33
6.5 Germplasm collecting and conservation 33
7 Genetic improvement strategies 36
8 Propagation 39
8.1 Pollen collecting, handling and controlled pollination 39
8.2 Seed 40
8.3 Asexual propagation 42
9 Agronomy of fruit and heart-of-palm production 44
9.1 Fruit: planting density and pruning 44
9.2 Fruit: mineral nutrition 44
9.3 Fruit: harvest and post-harvest handling 45
9.4 Heart-of-palm: planting density and pruning 46
9.5 Heart-of-palm: mineral nutrition 49
9.6 Heart-of-palm: harvest and post-harvest handling 52
9.7 Pests and pathogens 54
10 Production areas and commercial potential 58
10.1 Fruit 58
10.2 Heart-of-palm 59
11 Limitations and prospects of the crops 61
12 Future research needs 63
References 66

The Guilielma taxon (here considered a sub-genus of Bactris) probably originated in northwestern South America. By the Pleistocene species of this taxon were distributed along the Andes foothills from Bolivia to Panama. One of these species, or hybrids between several of them, gave rise to the pejibaye, which was then domesticated by the Amerindians. During the course of centuries, the pejibaye has become the most domesticated palm in the Americas, as attested by the great diversity of names, uses and fruit sizes. The principal use was of the starchy fruit, cooked for direct human consumption, fermented to make chicha, or ground and dried to make flour. Considerable genetic diversity and different Amerindian preferences have given rise to many distinct pejibaye races, varying in fruit size, shape and composition. A summary of what is known about the Amazonian races is presented. Potential modern uses are directed at further exploitation of some Amerindian uses and refinements of these: (1) heart of palm or palmito, and fruit (2) for direct human consumption, (3) for animal ration, (4) for flour and meal, and (5) for oil. The economic potential of each of these is mentioned. The breeding programs in Costa Rica and Brazil are outlined and preliminary plant ideotypes for each breeding objective are presented. The pejibaye has high immediate potential and will be even more useful once the breeding programs now underway are producing selected planting material. International support for these research programs is required however.

Neste trabalho estão apresentados os resultados da durabilidade natural do estipe (madeira) de Bactris gasipaes Kunth (pupunha), quando submetido ao ataque de insetos xilófagos, em ensaios em ambiente florestal e urbano. Foram utilizados dez palmeiras, cinco com espinhos e cinco sem espinhos, de plantios da Fazenda Experimental da Universidade Federal do Amazonas, localizada no km 40 da rodovia Manaus-Boa Vista (BR 174). De cada uma das palmeiras foram cortados três discos de aproximadamente 30 cm de espessura, retirados da base, do meio e do topo. No ambiente florestal, os discos foram distribuídos aleatoriamente, em área próxima ao plantio, no espaçamento de 0,5m, permanecendo durante 18 meses, período no qual foram efetuadas seis inspeções trimestrais para avaliar o grau de deterioração e coleta de insetos. Para o ensaio em condição urbana, os discos foram secionados axialmente para a retirada da medula e distribuídos aleatoriamente, nas posições côncava e convexa, sobre uma estr...

Commercial plantations of pejibaye (Bactris gasipaes Kunth, Palmae) are expanding in tropical America to supply the heart-of-palm market. The heart-of-palm is a gourmet vegetable composed of the tender unexpanded leaves in the palm's crown. Both spiny and spineless germplasm is available to growers, but there are no clear data on the superiority of either. Comparisons made among populations, among progenies within populations or among alleles within progenies are of interest for different reasons. In Hawaii, the Yurimaguas population (93% spineless; Pampa Hermosa landrace) is superior in heart-of-palm weight at one site and the Benjamin Constant (BC) population (79% spineless; Putumayo landrace) at another. Within the BC population there is a tendency for spineless plants to be superior in heart weight at one site and spiny to be superior at another. Within BC progenies, spineless plants often have more off-shoots at both sites, but superiority of spineless or spiny in terms of relative growth rates, heart weight and total edible weight depends upon location and seldom exceeds 10%. Although these yield components have low heritabilities there is variation amenable to selection, so that selecting within spineless germplasm should provide gains similar to those for spiny germplasm. Which is better is not a biological question but must be answered based upon comparative costs (management of spiny plants costs marginally to much more) and availability of germplasm at a given location.

The peach palm (Bactris gasipaes Kunth) is the only Neotropical palm domesticated by Native Americans. Its place of origin as a crop (B. gasipaes var. gasipaes) has been debated for more than a century, with three hypotheses currently in discussion: southwestern Amazonia; northwestern South America; or multiple origins in the distribution of the wild relatives (B. gasipaes var. chichagui). The small amount of archaeological data available supports the second hypothesis, but they contrast dramatically with the molecular-genetic analyses that support the first or the third, depending on how they are interpreted. On morphological grounds, two of the three types of var. chichagui are plausible candidates for wild ancestral populations. All the molecular-genetic analyses have identified a deep division between the landraces of cultivated peach palm in western Amazonia to Central America and those in southwestern to eastern Amazonia. The first analysis using isoenzymes linked the Tembe population (Bolivia) with the Pará landrace (eastern Amazonia), and these were distant from the western landraces. Multiple RAPD and SSR analyses identified the same deep division, which was interpreted by the group of researchers in Brazil as a single domestication in southwestern Amazonia with two dispersals, while another group working in Costa Rica interpreted it as three domestication events. Analysis with nuclear markers does not allow discrimination among the hypotheses, because gene flow may occur via pollen and seed. A new analysis with two sequences from the chloroplast genome, which has maternal inheritance and is therefore more appropriate to test the hypothesis, suggests that the cultivated peach palm was domesticated once in southwestern Amazonia, with two dispersals. One dispersal started in the upper Ucayali River basin, in southeastern Peru, and then throughout western Amazonia, northwestern South America and southern Central America. Another dispersal started in the upper Madeira River basin and then along the Madeira River into eastern Amazonia. New explorations in southwestern Amazonia are critical to identify the exact location of the original events.

A review of palm remains recorded at archaeological sites throughout the New World is presented. Remains have been found at 130 sites from the southern United States to southern Uruguay. They are of four kinds: carbonized or dry endocarps or seeds, phytoliths, pollen, and implements. Twenty-nine genera and at least 50 species of palms (i.e., about 9% of all American species) have been recorded. The oldest record dates back to 14,700b.p. for carbonized endocarp fragments of an unidentified palm in Rondônia, Brazil. The use of palms, as recorded from remains, was particularly widespread after 9000b.p. The predominant remains are endocarps ofAcrocomia, Attalea s.l.,Astrocaryum, Bactris, Syagrus, Elaeis, andOenocarpus, all of which are important sources of edible oils or edible fruits and are still widely used by aboriginal peoples. The review supports the hypothesis that human groups have played an important role in the dispersal of some palm species in the neotropics. Humanaided dispersal ofAcrocomia aculeata from South America into Central America, and ofOenocarpus bataua from northwestern Amazonia to other areas, is postulated. Archaeological remains support the hypothesis that pejibaye (Bactris gasipaes) was domesticated in the inter-Andean valleys or on the adjacent Pacific lowlands of Colombia and later introduced into the Amazon Basin. Se presenta una revisión de los registres de vestigios de palmas en yacimientos arqueológicos en el Nuevo Mundo. Se han encontrado vestigios en 130 yacimientos, desde el sur de los Estados Unidos hasta el sur de Uruguay. Los vestigios son de cuatro tipos: endocarpos quemados o secos, fitolitos, polen e implementos. Representan 29 géneros y por lo menos 50 especies de palmas (es decir, cerca del 9% de todas las especies americanas). El registro más antiguo data de 14,700a.p., para fragmentos de endocarpo carbonizados de una palma no identificada en Rondônia, Brasil. El uso de las palmas, tal como lo evidencian los vestigios, estuvo particularmente extendido después del 9000a.p. Los vestigios predominantes son endocarpos de los génerosAcrocomia, Attalea, Astrocaryum, Bactris, Syagrus, Elaeis, yOenocarpus, los cuales son fuentes importantes de aceites o frutos comestibles, todavía usados por pueblos aborígenes. La revision apoya las hipótesis según las cuales los grupos humanos han jugado un importante papel en la dispersión de algunas palmas en el neotrópico. Se postula la dispersión antropogénica deAcrocomia aculeata desde Suramérica hacia Centroamérica y deOenocarpus bataua desde el noroeste de la Amazonia hacia otras áreas. Los vestigios arqueológicos apoyan la hipótesis de que el chontaduro (Bactris gasipaes) fue domesticado en los valles interandinos o en las tierras bajas aledañas del Pacífico en Colombia, y después introducido a la Amazonia.

RESUMO Embora a pupunha possa ser propagada de duas formas, por via seminal ou por via vegetativa, na prática a propagação é quase exclusivamente por sementes. A abundância de relatos entre populações tradicio-nais sugere, entretanto, que a propagação vegetativa foi usada ocasionalmente, e ainda hoje os povos tradicionais afirmam praticar esta técnica. Em face dessas observações antagônicas, decidiu-se confirmar a veracidade das declarações dos agricultores de duas comunidades caboclas da Amazônia Central com o estudo do DNA usando a técnica molecular RAPD (polimorfismo de DNA amplificado ao acaso). Nessa análise quinze amostras de pupunha foram coletadas junto a quatro produtores do Estado de Amazonas, sendo quatro amostras de dois sítios do Ramal Nova Esperança, Manacapuru e onze amostras de dois sítios da Estrada de Autazes. Os quatro iniciadores geraram 48 produtos de amplificação úteis (bandas), com uma média de 12 bandas por iniciador. As similaridades de Jaccard mostrou um valor de 0,808. O dendrograma baseado nas similaridades de Jaccard e Distância Genética de Nei mostraram dois grandes grupos. Esses grupos se dispuseram a uma distância de 0.55, que é uma distância esperada entre espécies diferentes e não entre raças dentro de uma espécie. Os demais indivíduos formaram subgrupos sem apre-sentar ordem aparente. Palavras-chave: Análise Genética. Distâncias Genéticas. Relaões Filogenéticas. Conhecimento Tradi-cional ABSTRACT Although the peach palm can be spread in two ways, for seminal road or for vegetative road, in the practice the propagation is almost exclusively for seeds. The abundance of reports among traditional populations suggests, however, that the vegetative propagation was used occasionally, and still the traditional people affirm today to practice this technique. In face of those antagonistic observations, decided to confirm the truthfulness of the farmers of two community's caboclas of Central Amazônia declarations with the study of the DNA using the molecular technique RAPD (Random Amplified Polymorphic DNA). In that analysis fifteen peach palm samples they were collected four close to producing of the State of Ama-zonas, being four samples of two ranches of the Extension New Hope, Manacapuru and eleven samples of two ranches of the Highway of Autazes. The four initiators generated 48 useful amplification products (bands), with an average of 12 bands for initiator. The similarities of Jaccard showed a value of 0,808. The dendrograma based on the similarities of Jaccard and Genetic Distance of Nei showed two great groups. Those groups were disposed at a distance of 0. 55, that is a distance waited among different species and don't enter races inside of a species. The other individuals formed subgrups without presenting apparent order.

Abstract
The percent composition, soluble and insoluble food fibers, oil fatty acids and minerals were determined in the mesocarp of fruits of three peach palm (Bactris gasipaes Kunth) populations grown in Central Amazonia, Brazil. Amino acids were also determined in one of the populations. The mean protein levels ranged from 1.8 to 2.7%, lipid levels ranged from 3.5 to 11.1%, the nitrogen free fraction ranged from 24.3 to 35%, food fiber ranged from 5.2% to 8.7%, and energy ranged from 179.1 to 207.4 kcal%. All essential, as well as non-essential, amino acids were present, with tryptophan and methionine presenting the lowest mean concentrations. The mono-unsaturated oleic acid predominated in the oil, ranging from 42.8 to 60.8%, and palmitic acid was the most abundant saturated fatty acid, ranging from 24.1 to 42.3%. Among the essential fatty acids, linoleic acid was the most abundant, with a maximum of 5.4% in Pampa-8. The most important mineral elements were potassium, selenium and chromium, respectively corresponding to 12%, 9% and 9% of daily recommended allowances. Considering the nutritional potential of the fruit, we suggest its more frequent incorporation into the diet of the Amazonian population.

Various factors affect the induction of somatic embryogenesis in peach palm (Bactris gasipaes Kunth). Among these, both the type and level of auxins had the greatest influence on in vitro responses, although the genotype and the developmental stage of the explants also influenced results. Younger inflorescences were more competent to respond to SE induction than more mature inflorescences and the use of a pre-treatment with 2,4-D (200 μM) in liquid MS culture medium also increased the embryogenic capacity, and diminished the development of flower buds. Higher oxidation rates were observed in explants maintained on 2,4-D-supplemented culture medium, while on 300 μM or 600 μM Picloram and Dicamba lower oxidation rates were observed. The progression from floral meristem to flower bud occurred at high frequency when low concentrations of auxins were used, independent of the type. Higher concentrations of Picloram or Dicamba reduced or even inhibited flower bud development. Picloram also...

Peach palm (Bactris gasipaes) is cultivated by many indigenous and traditional communities from Amazonia to Central America for its edible fruits, and is currently important for its heart-of-palm. The objective of this study was to investigate the mating system of peach palm, as this is important for conservation and breeding. Eight microsatellite loci were used to genotype 24 open-pollinated progenies from three populations of the Pampa Hermosa landrace maintained in a progeny trial for genetic improvement. Both the multi-locus outcrossing rates (0.95 to 0.99) and the progeny level multi-locus outcrossing rates (0.9 to 1.0) were high, indicating that peach palm is predominantly allogamous. The outcrossing rates among relatives were significantly different from zero (0.101 to 0.202), providing evidence for considerable biparental inbreeding within populations, probably due to farmers planting seeds of a small number of open-pollinated progenies in the same plot. The correlations of paternity estimates were low (0.051 to 0.112), suggesting a large number of pollen sources (9 to 20) participating in pollination of individual fruit bunches. Effective population size estimates suggest that current germplasm collections are insufficient for long-term ex situ conservation. As with most underutilized crops, on farm conservation is the most important component of an integrated conservation strategy.

Allozyme variation was examined in three pejtbaye (Bactris gasipaes) populations introduced into Hawaii for use in heart-of-palm improvement: Benjamin Constant (Putumayo landrace), San Carlos (Guataso landrace) and Yurimaguas (Pampa Hermosa landrace). Nine enzymes encoded by 16 putative loci with a total of 38 alleles were resolved from meristem extract. Five loci were fixed in all populations, four additional loci were fixed in SC and one additional locus in BC. Six of the 38 alleles were rare, with three unique to BC and one to Y; two moderately high frequency alleles were unique to SC. Mean number of alleles per locus was lowest (1.6) in SC, with BC and Y having 2.1 alleles. Percentage of polymorphic loci was lowest in SC (44) and highest in Y (69). Observed heterozygosity was lowest in SC (0.051) and BC (0.066) and highest in Y (0.141). Nei's genetic identity was 0.985 between BC and Y and averaged 0.952 between SC and BC-Y. The low heterozygosities are probably due to a long history of selection and inbreeding (sib-mating) during the domestication process, followed by intensive recent selection for spinelessness and more inbreeding. All populations are very closely related, suggesting a single domestication event in Amazonia.

Os conhecimentos de nutrição mineral da pupunheira (Bactris gasipaes Kunth, Palmae) são relativamente escassos e incipientes, faltando dados consistentes sobre sua demanda nutricional desde a fase de viveiro até a fase de produção. Com o objetivo de caracterizar os sintomas de carências de macro e micronutrientes, e identificar os níveis analíticos associados às carências minerais em mudas de pupunheira, foi conduzido um ensaio em casa de vegetação, pertencente ao Instituto Nacional de Pesquisas da Amazônia, Manaus, AM. Plantas de pupunheira foram cultivadas em solução nutritiva, tendo como substrato areia lavada. Os tratamentos foram: completo e omissão individual de nitrogênio, fósforo, potássio, cálcio, magnésio, enxofre, ferro, boro, cobre, manganês, zinco e molibdênio. O delineamento experimental foi inteiramente casualizado com treze tratamentos e quatro repetições. As deficiências dos nutrientes nas mudas de pupunheiras se traduziram por sintomas típicos e facilmente identifi...