When was the papaya domesticated

Under should be made to know and preserve this important species cultivation papaya trees grow fast, producing mature fruits within in its wild form. Moreover, because papaya is one of the most 9—12 months after planting.

Commercially, a density of 1,— economically important tropical crops in the world, and wild 2, trees per hectare, can produce from , to , lbs populations still occur naturally, the species represents an ideal per hectare, per year Gonsalves, Among common fruits, system to study, in depth, the process of domestication.

Here, papaya is ranked first on nutritional scores for the percentage we present a review of the existent knowledge on the origin, of vitamin A, vitamin C, potassium, folate, niacin, thiamine, distribution and domestication of papaya, and discuss the usage riboflavin, iron and calcium, and fiber Huerta-Ocampo et al.

Moreover, fruits, stems, leaves and roots of papaya are used this valuable species. Because of its rapid growth, continuous harvest and Different authors suggest a Mesoamerican origin of C.

B Wild population of Carica papaya in Yucatan, Mexico. C Wild fruits, intermediate form fruits possible hybrid between wild and domesticated plants , and domesticated papaya fruit Maradol variety. One reason for the chromosome in cultivated hermaphrodite individuals of papaya lack of archaeological data is the difficulty to identify papaya VanBuren et al. By sequencing the entire male-specific from phytoliths, and their pollen grains have been hardly region of the Y chromosome and comparing it with the found Carvalho and Renner, Together, the presence of previous sequences of the hermaphrodite-specific region of the undomesticated populations in Mexico and Central America and Yh chromosome, Wang et al.

Carvalho and Renner Given the ubiquity of dioecy in wild populations of papaya obtained a molecular phylogeny of the Caricaceae family using in Mesoamerica, an origin for the species in this region is chloroplast and nuclear data 4, bp of the 34 species in the a more parsimonious hypothesis.

Yet, genomic studies can family. This is the only phylogeny using members from all the aid to determine the origin of some innovations related with Caricaceae family. Their resulting phylogeny points that Carica domestication. Additionally, thought Carvalho and Renner, Moreover, Carica, Jarilla domestication is expected to reduce genetic diversity and and Horovitzia show an unilocular ovary, whereas the remaining provoke selective sweeps in genes associated to characters target South American Caricaceae possess 5-locular ovaries Carvalho of domestication Purugganan and Fuller, A recent and Renner, This morphological synapomorphy supports phylogeographic analysis of 19 wild populations of papaya in the family phylogeny.

Mexico using nuclear and chloroplast markers, revealed high Frontiers in Ecology and Evolution www. The higher genetic diversity was found in locations of Indies and tropical Asia Storey et al. After that, it rapidly southern Mexico, suggesting this region as a genetic reservoir for spread into other Asian countries and finally to Africa, brought the species see below.

Summing up, evidence strongly suggests by European colonial powers, such as Portugal, Denmark, Great that C. Moreover, because the papaya seeds have a moderate period Regarding the natural distribution of papaya, this has been of longevity, it is likely that this trait aided to its rapid spread suggested to range from the northern tropical limit of Mexico to throughout the tropics, where it has existed practically since man Costa Rica in Central America Aradhya et al. Nowadays, there and Renner, However, the precise assessment of its natural are many varieties of papaya cultivated in most tropical and distribution is still lacking.

One reason for this is the scarcity subtropical regions of the world, differing in traits such as fruit of studies in wild populations of papaya, and the paucity of size, color, flavour, and tree size Moore, In general, they differ the distribution of papaya. The principal characters of papaya that have been studied wild papaya is high enough as to endanger the persistence of as target of selection under domestication are: tree size, fruit size, plant species Barlow et al.

This, coupled with the lack sex types, and morphology, and germinability of seeds. Whereas of information on the state of many wild varieties of important tree size has been selected to become smaller to facilitate fruit crop species, warn us about the relevance of studying and harvesting Niklas and Marler, , selection has been focused conserving wild populations and wild relatives of papaya and on enlarging the fruit size and increasing the ovary wall pulp other crop species.

Regarding sex types, in dioecious fruit crops, mutations The limited occurrence of wild populations of C. In the Caricaceae family, region Carvalho and Renner, The Maya was the most important mutation in male plants and were likely selected by humans culture present in that region before the conquest of Mexico for its favorable fruit phenotype Ueno et al.

Moreover, by Spain in the sixteenth century, and probably one of the the region of the chromosome that produces hermaphrodite first to cultivate and trade the fruits of C.

Although crop plant Mesoamerican people to tenderize meat by wrapping it in papaya domestication in Mesoamerica occurred around 6, years ago, leaves and that this knowledge was then taken to Europe after the the estimation coincides with the rise of the Maya civilization. By the time of the Given that hermaphrodite individuals are rarely found in conquest of Mexico, it is believed that papaya was cultivated by wild populations of Mesoamerica, this strongly supports that native people all the way from southern Mexico to the Isthmus papaya was domesticated by the Mayans or other Mesoamerican of Panama, where it was locally known as olocoton Storey et al.

It is believed that the rare allele It was introduced into the Hispaniola island nowadays Haiti and Santo Domingo in 1, Finally, seeds from wild and domesticated papayas differ in There, it acquired the Carib Indian name ababai, that would size, germination rate, dormancy, and light sensitivity. Paz and then be changed to papaia, papia, papeya, and finally papaya. In papaya was taken outside of America to lessening the importance of specific environmental conditions Frontiers in Ecology and Evolution www.

This domestication Tripathi et al. For genetic diversity studies, some syndrome has been reported for other crops Doebley et al. Ocampo et al. Understanding crop domestication is crucial to fulfill the Our recent surveys about the genetic diversity and structure of demand for improving yield and quality of crops Tang et al.

Using both nuclear SSRs for the conservation of their genetic resources, and effective and chloroplast DNA markers in individuals of 19 natural crop improvement Moore, Furthermore, crops represent populations, we found a mean observed heterozygosity of 0. For the case of range of 0. The area with the higher genetic diversity for both to have its genome sequenced Ming et al.

With the chloroplast DNA, we found a At the local scale, we state of wild populations, to recent advances in domesticated also found a negative effect of habitat fragmentation for wild varieties.

Ming et al. Populations that inhabit forest fragments showed a reduced A recurrent consequence of domestication is the reduction genetic diversity, higher population differentiation, and less of genetic diversity due to the 2-fold effect of genetic drift migrants. Together, these studies suggest that wild papaya has and selection that operate during the domestication process maintained genetic connectivity among populations throughout Doebley et al.

However, concerns for the species in its wild form. Genetic diversity of cultivated and feral papayas has shown Early on, the use of genetics in tropical fruit crops was to be, in general, lower than wild populations: 1 Using AFLPs confined to the development of isozyme and dominant PCR- markers, a genetic similarity of 0.

Since then, codominant markers, known as simple He of 0. Years of selective breeding would explain the leading to increased interest in tropical tree fruit crops and low genetic diversity in the domesticated papayas brought about to new opportunities to increase the rate of genetic gain in the domestication bottleneck.

Recent approaches During domestication, gene flow between wild and in massive sequencing tools, such as de novo genome sequencing, domesticated conspecific plants plays an important role in whole genome resequencing, reduction of genome complexity addition to other evolutionary forces such as genetic drift and using restriction enzymes, transcriptome analysis, and epigenetic selection. Anaya et al. However, there are no studies evaluating the and Wing, Further efforts were made to complete the ecological and evolutionary consequences of this phenomenon.

The papaya genome has shown to be three times the consideration. Amplifications in have been released for commercial production in Hawaii, USA , gene number with roles in the evolution of tree-like habit, in by public institutions in the USA Tecson Mendoza deposition and remobilization of starch reserves, attraction of et al. This transgenic variety was seed dispersal agents, and adaptation to tropical day lengths have developed to control the papaya ringspot virus PRSV. The been identified Ming et al.

After a spread of the virus in that nearly killed Recent genetic and genomic methods have revealed all cultivars in Puna Island in Hawaii, efforts to develop a interesting facts about domestication in papaya.

For instance, resistant variety to the PRSV began. Two varieties successfully Wu et al. Analysing the in Hawaii: Rainbow and SunUp. However, some concern raised regarding the potential found evidence of a strong selective sweep, as genetic diversity effects of gene flow between transgenic and nontransgenic showed a reduction at the recessive y allele in comparison with cultivars.

Experiments were carried out to assess this; from the dominant Y allele present in the yellow-fleshed papayas through , Gonsalves et al.

Although cultivars of transgenic papaya are confined feral intermediates in Costa Rica Wu et al. In other to Hawaii, little is known about the ecological, evolutionary study, Porter et al. This group of genes are the most common in plant Silva-Rosales et al. Nowadays, transgenic papaya from disease resistance proteins. Given that PRSV is widespread Furthermore, papaya has been recognized as an excellent in papaya cultivars around the world, papaya SunUp could model for studies about sex determination in plants Ming serve as a transgenic germplasm source to breed suitable virus- et al.

However, despite these Weingartner and Moore, , origin and evolution of dioecy achievements, many countries have rejected transgenic crops; in in the Caricaceae family, and the identification of candidate genes , only transgenic papaya and plum had been approved for and genome-wide DNA markers for papaya improvement Ming human consumption Litz and Padilla, Because the fruit is also a good source of fiber, papaya isn't only physically satisfying — it'll also help you stay full longer.

As a result, you may end up consuming fewer calories throughout the day. The carrot Daucus carota subsp. They are a domesticated form of the wild carrot , Daucus carota, native to Europe and Southwestern Asia.

The carrot is a biennial plant in the umbellifer family Apiaceae. Papaya : Natural antioxidants within the fruit make papaya a great choice for diabetics. Diabetics are prone to many ailments, including heart or nerve damage caused by irregular blood sugar levels.

Being a diabetic should never stop you from eating fruits. Carica papaya L. Papaya is a tall herbaceous plant in the genus Carica; its edible fruit is also called papaya. It is native to the tropical region of America, mainly from southern Mexico to Central America. Now these plants are grown in all tropical regions of the world. Papayas also contain healthy antioxidants known as carotenoids — particularly one type called lycopene.

Summary The papaya is a tropical fruit high in vitamins C and A, as well as fiber and healthy plant compounds. It also contains an enzyme called papain, used to tenderize meat. For maximum results, people looking to lose weight quickly should consume papaya during breakfast and as a snack, between lunch and dinner. For breakfast, pair papayas with a source of good quality protein and a small amount of healthy fats. Eating papaya as a healthy snack after lunch, helps you stay full for longer.

Benefits of Eating Papaya for the Skin The naturally high vitamin A and papain content found in papaya helps break down the proteins and gives your skin elasticity and increases your collagen levels. Papaya contains lots of water that hydrates the skin from inside out, leaving your skin moisturized and smooth. Papaya may cause severe allergic reactions in sensitive people. Papaya latex can be a severe irritant and vesicant on skin.

Papaya juice and papaya seeds are unlikely to cause adverse effects when taken orally; however, papaya leaves at high doses may cause stomach irritation. Kiwi berries are edible berry - or grape-sized fruits similar to fuzzy kiwifruit in taste and appearance, but with thin, smooth green skin. They are primarily produced by three species: Actinidia arguta hardy kiwi , A. A pineapple is neither a pine nor an apple, but a fruit consisting of many berries that have grown together.

This also means that Pineapples are not a single fruit, but a group of berries that have fused together. The scientific name of a pineapple is Ananas comosus. Choose a Ripe Papaya You know if you have ripe papaya if it has skin that is turning from green to yellow. If papaya is ripe , you should be able to press your thumb into the flesh.

Papayas will ripen more quickly when put in a paper bag with ethylene-producing fruits, such as apples or bananas. The PRSV acts destroying the photosynthetic capacity of the leaves, leading first to a decrease in fruit quality and yield, loss of vegetative vigor, and eventual death of plants Lius et al.

After a spread of the virus in that nearly killed all cultivars in Puna Island in Hawaii, efforts to develop a resistant variety to the PRSV began. Two varieties successfully controlled the virus infection and saved the papaya production in Hawaii: Rainbow and SunUp. However, some concern raised regarding the potential effects of gene flow between transgenic and nontransgenic cultivars. Experiments were carried out to assess this; from through , Gonsalves et al.

Although cultivars of transgenic papaya are confined to Hawaii, little is known about the ecological, evolutionary and commercial consequences of possible transgene flow in the center of origin of papaya where natural populations occur, and where attempts to release transgenic cultivars have prevailed Silva-Rosales et al. Nowadays, transgenic papaya from Hawaii is consumed in USA, Canada, and Japan, but no studies have been made to assess the possible movement of seeds to papaya's origin center.

Because transgenic papaya saved Hawaii's papaya industry, it has been recognized as a model species of translational biotechnology and the best-characterized commercial transgenic crop. Given that PRSV is widespread in papaya cultivars around the world, papaya SunUp could serve as a transgenic germplasm source to breed suitable virus-resistant cultivars Ming et al.

However, despite these achievements, many countries have rejected transgenic crops; in , only transgenic papaya and plum had been approved for human consumption Litz and Padilla, With the recent advances of DNA next generation sequencing a great interest has been advocated to crops.

Papaya is the fifth flowering plant after Arabidopsis, rice, poplar, and grape to have its genome sequenced Kanchana-Udomkan et al. A draft genome approximately 3x coverage was obtained from a transgenic female cultivated papaya from the SunUp variety from Hawaii, using a whole-genome shotgun approach Ming et al. Further efforts were made to complete the draft genome of papaya; Yu et al. The papaya genome has shown to be three times the size of the Arabidopsis genome, containing fewer genes, with a minimal angiosperm gene set of 13, Amplifications in gene number with roles in the evolution of tree-like habit, deposition and remobilization of starch reserves, attraction of seed dispersal agents, and adaptation to tropical day lengths have been identified Ming et al.

Recent genetic and genomic methods have revealed interesting facts about domestication in papaya. For instance, Wu et al. Analysing the levels and patterns of genetic diversity at the CYC-b locus y allele and six loci in a kb region flanking this locus, they found evidence of a strong selective sweep, as genetic diversity showed a reduction at the recessive y allele in comparison with the dominant Y allele present in the yellow-fleshed papayas wild and cultivated.

By means of a haplotype network, the authors suggest that the y allele likely originated in the wild and was introduced into the domesticated varieties during the domestication of papaya.

However, they found shared haplotype structure among some wild, feral, and cultivated haplotypes near the y allele, suggesting a successive escape of the y allele from red cultivars back into wild populations, probably through feral intermediates in Costa Rica Wu et al.

In other study, Porter et al. This group of genes are the most common in plant disease resistance proteins. The authors argue that the absence of recent genome duplication and relatively low gene number in papaya, may explain the apparent scarcity NBS gene number, and that this could function as pathogen surveillance, making papaya suitable for functional studies and a better understanding of plant resistance at the genetic level Porter et al.

Furthermore, papaya has been recognized as an excellent model for studies about sex determination in plants Ming et al. These studies have helped to expand the knowledge for tropical fruit tree genomics. Moreover, phylogenetically, papaya belongs to the Brassicales, and can serve as an excellent out-group to study genome evolution in the Brassicaceae family, given that it shares a common ancestor with Arabidopsis , about 72 MYA Ming et al. Undoubtedly, the sequencing of the papaya genome has opened avenues to investigate the function of many genes for which little is known.

The sequenced genome will facilitate knowledge about the genomic regions associated to diverse aspects of plant defense, plant growth, development of leaves, roots, and fruits, flowering, as well as fruit ripening, and circadian clock Tripathi et al.

Regarding its commercial use, a variety of markers have been developed to distinguish between the different sexes in juvenile plants Somsri et al. This aided in eliminating females, given that hermaphrodites are preferred for their bigger and elongated fruits, rather than the rounded female fruits that require greater container space for shipping Tripathi et al.

However, in commercial production, males are useless and with these markers, males couldn't be distinguished. Recently, Liao et al. Given that sex determination is only possible around 6 months after germination, these markers largely facilitate the elimination of undesired plants.

In spite of these advances, many questions remain unsolved for papaya, mainly for the wild counterpart. Compared to annual crops and temperate trees, genetic improvement of tropical fruit trees has been limited Litz and Padilla, The use of genomics for the development of new tropical cultivars adapted to higher temperatures and increased droughts will become necessary to alleviate food security in the upcoming years Abberton et al.

The organization of genetic diversity in wild relatives represent important genetic resources that can facilitate the development of climate-tolerant cultivars Gepts, Finally, the continuous advances of genomic tools in crop species will also serve to gain insights into the ecological and evolutionary knowledge of domestication in tropical crops. For instance, the functional genomics aspects of domestication will help assess how many and which genes show differences in expression between wild and domesticated types Sarah et al.

Plant domestication represents one of the most relevant events in human history. Today, papaya is one of the most important economically crop species worldwide.

Understanding the ecology, evolutionary history and domestication process of such species is necessary to maintain food security in the future and counteract upcoming threats of overpopulation and climate change.

Papaya represents the third most produced crop in the tropics worldwide and an important source of commercial uses for humans. It was most probably originated and domesticated in Mesoamerica where wild populations still occur and distribute in Mexico and Central America.

In nature, wild papayas play an important role in the regeneration of their natural habitat and possess high levels of genetic variation, representing the genetic wealth and evolutionary potential of the species.

Finally, papaya is an excellent model for genomic studies as it is one of the first plant species to have its complete genome sequenced. However, there is still much more knowledge needed from this important species, mainly about its wild populations and the evolutionary process of domestication.

MC-P wrote the manuscript. JN-F reviewed drafts of the manuscript and contributed to writing the final version. Both authors were involved in the final editing and review of the paper.

The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest. The handling Editor declared a shared affiliation, though no other collaboration, with one of the authors JN and states that the process nevertheless met the standards of a fair and objective review. Abberton, M. Global agricultural intensification during climate change: a role for genomics.

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