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Cherimoya reproductive morphology

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Male stage Shiva Nia
Contents of this annotation:
1. Cherimoya reproductive morphology
female stage Shiva Nia
Contents of this annotation:
1. Cherimoya reproductive morphology
female stage Shiva Nia
Contents of this annotation:
1. Cherimoya reproductive morphology

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Annona cherimola: The most delicious fruit known to man Cherimoya is a fruit from the genus Annona, it is cone shaped, green, leathery skin with a creamy and sweet flesh. The fruit has a sweet taste and it is rich in nutrients and minerals. Due to its creamy textures, it is also called the custard apple. It is believed to be native to the Andes Mountain range in South America but it has been distributed and planted in different areas of the world today. One of the major problems for cherimoya cultivation is inadequate natural pollination since the male and the female structures of each flower do not mature at the same time. Due to its rich mineral content cherimoya is known to have some health benefits including lowering blood pressure. However, cherimoya and other Annona species contain annonacin which is a toxin that can affect the nervous system and the brain. Some studies have suggested that high amounts of consumption of Annona fruits can lead to increased risk of a specific type of Parkinson’s disease that does not respond well to medication. The seeds can be toxic, so when eating the fruit, it is best for the skin and seeds to be discarded. While Cherimoya has lots of health benefits and can be delicious it needs to be eaten in moderation to avoid its negative effects on the nervous system. by Shiva Nia, Kyi Ther Min #Cherimoya Nomenclatural History
Annonaceae cherimola Miller or otherwise known by the common names of chirimoya, cherimola, or cherimoya.The common name of cherimoya originated from the Quechua word chirimuya which means “cold seeds”. This is because the tree itself only grows at high altitudes with colder temperatures and the seeds will only germinate in those conditions (Morton 1987). However, the creamy texture of the flesh and aroma also give it its secondary common name of the “custard apple”.
Other names used for cherimoya in different countries:
In Venezuela, it is called chirimorrinon
In Brazil, graveola, graviola, or grabiola
In Mexico, pox or poox; in Belize, tukib
In El Salvador it is sometimes known as anona poshte; and elsewhere merely as anona, or anona blanca.
In France, it is anone; in Haiti, cachiman la Chine
Indian names in Guatemala include pac, pap, tsummy and tzumux
Evolutionary History
One of the earliest evolutionary branches in the angiosperm phylogenetic tree is the magnoliid clade. According to APG IV, the Magnoliids clade consists of about 10,000 species, 18 families, and four orders. The orders are Magnoliales, Canellales, Laurales, and Piperales. Magnoliales order is a taxonomic group of woody or herbaceous flowering plants which consists of about 3,000 species, 154 genera, and six families. The families in the order are Annonaceae, Myristicaceae, Magnoliaceae, Degeneriaceae, Eupomatiaceae, and Himantandraceae. Annona cherimola is a part of the larger Annonaceae family, which contains about 130 genera and 2400 species as well as being one of the oldest families among the flowering plants with an ancestry of roughly 95 million years. Among the genus Annona, which diverged 25 million years ago, there are around 175 species. Some common species under this genus include soursop, sugar apple, and atemoya which is a hybrid cross between cherimoya and sugar apple. Molecular data suggests that the A. cherimoya and the A. squamosa (sugar apple) are the closest relatives amongst the fruiting species (Rare Fruit Club, 2021).
Cherimoya genetic diversity is highest, in the countries of its origin, Peru and Ecuador. Whereas, in Spain and Madeira it has less genetic diversity which can be due to genetic bottlenecks effect from limited introduction. Cherimoya fruits’ genetic diversity is also high in california which is possibly due to the expansion of cherimoya cultivation in this region (Zonneveld et al. 2012).

Ecology and Life History
The cherimoya tree is a subtropical or mild-temperate tree that is sensitive to climate. It requires long days and is typically grown in high elevations between 4,000 and 8,000 ft where the temperature ranges from 62.° and 77°F (Morton, 1987). Any temperatures lower than 26°F can severely kill or injure the mature tree. It also prefers drier and non-windy climates for proper pollination. The seeds, if properly stored, are viable for several years. To properly plant, soak in water for 1 to 4 days and discard those that float. Plant directly in the soil and will germinate in 3 to 5 weeks (Morton, 1987). Once the plants are 3 to 4 inches high they are transplanted to nursery pots and within a year to two they bud and are grafted, growing up to 3 to 4ft (Morton, 1987). To properly grow the tree, it is recommended to interplant with maize or potatoes to provide some organic material to the soil it grows in. The tree goes dormant during the winter for around 4 months and once the dormancy period is over, leafbuds begin to appear and yield fruits around 3.5 to 5 years old. Production ramps up during the 5th to 10th year where there is an expected yield of 25 fruits per tree. The ideal time to pick the fruits is when its full grown but firm as ripening is best off the tree. There should be a slight yellow-greenish hint on the skin. In Bolivia, they shake the fruit to listen to the sounds of loose seeds to judge if its ready to be picked. Once picked, the fruit should be stored at room temperature and will soften and be ready to eat in 3 to 4 days. The cherimoya tree also doesn’t rely on any pollinators that are known and heavily require wind or hand pollination to successfully reproduce. The only hypothesized pollinator for the cherimoya is a small beetle native to the region that are able to spread the pollen from the fleshy part of the female flower and male flowers, whereas bees are too large to get too both.

Vegetative Morphology
The tree mostly remains green and functional through different seasons, it is a relatively fast-growing tree with low-branched and spreading tree. However , the roots are slow growing and weak when the tree is young. It develpoved a taproot system initially, which them become fibrous roots that will stay shallow unless deep watered. This plant is often grafted or budded onto seedling rootstock.They have a grey trunk with relatively smooth bark. The height can range from 16 to 30 ft tall. The mature branches are woody and sappy. The leaves are leathery and are approximately 2 to 9.8 inches long and 1.2 to 3.9 inches wide. The leaves are briefly deciduous to semi-deciduous. They have an elliptic shape, and they are more rounded near the leaf stalk with more pointed ends. The leaves are single and they alternate. They usually have a darker green color and they are slightly hairy on the upper surface and slightly furry on the underside (Cautín and Agustí 2005).

Reproductive Morphology
The flowers are pale green and they are about 1.2 inches long. They are dichogamous and borne in either solitary or groups of 2-3 on short stalks. The flowers themselves have three oblong petals that have a slightly pink and white interior and have a green or brown exterior. They are fragrant and have a strong fruity odor.While the plant is hermaphroditic which means they have both male and female reproductive organs they have a mechanism that avoids self-pollination. The mechanism that prevents cherimoya from self pollination happens because of the dichogamous characteristic of this plant which means that the flowers have the stamen and pistil mature at different stages (Richardson and Anderson 1996).
The first reproductive stage of cherimoya is the female stage. The flower’s sticky stigmas can receive pollen only from another flower and receptivity reduces in arid mountain air. The weather condition and temperature can affect the female flower’s receptivity period, for example humidity can increase it. The second stage, male stage, happens one to three days after the female stage. At this stage petal separates all the way and anther produces pollen so pollinators distribute it far and wide to other flowers or gets distributed through air. The cherimoya’s natural pollinator is a tiny beetle. These natural pollinators do not live in California therefore one of the major problems for cherimoya pollination is inadequate natural pollination. One solution to this is the Hand-pollination which is more common when growing fruits (Schroeder 1943).
The fruit is edible and the appearance is conical, almost heart-shaped. The interesting skin of the fruit is formed from the fusion of the carpels which results in the bumpy texture. They are usually 4 to 8 inches long and 4 inches in width. They weigh around 18 ounces. The fruit can be easily broken or cut open. The inside is white, juicy and creamy with a pleasing aroma. The black bean-like seeds are approximately ½ to ¾ in length and scattered throughout the flesh once opened(Cautín and Agustí 2005).

Historical Accounts
There is a historical account that the seeds of the cherimoya were sent by P. Bernabe Cobo from Guatemala in 1629 and thirteen years after the cherimoya fruits and trees were found in markets and cultivation in Lima. Cobo described the fruits as “[…] white fleshed and very smooth, with a sweet and sour taste […] the best and cheapest of all the fruit of the Indies […]” (Museo Chileno de Arte Precolombino, n.d). Cobo also sent the seeds to Mexico in 1629, where, now, it thrives in the higher elevations. In 1757, it was introduced to Spain as a dooryard tree but in the 1940s, the cherimoya tree became a staple and replacement for the orange trees that were wiped out due a disease. Today, Spain is one of the biggest producers of cherimoya. In 1871, the seeds from Mexico were brought over to California and planted. 9,000 trees were plants in 1936 but were killed off by too cold of a climate in 1937 and only small orchards remain. California remains the only state in the United States that can successfully cultivate cherimoya and bear fruits (Morton, 1987).
The cherimoya has been used and seen in ceramic art from the Inca during the Late Horizon period around 1493. This archaeological find of the ceramic vessel is on display in the Chimu-Inca style collections of the Musee de l'Homme in Paris that was donated by M. Lencoyne in 1887. The Incan people historically used cherimoya as a dessert and the sweet custardy flesh was a delicacy to many and Chileans consider the cherimoya to be their “national fruit” (National Research Council, 1989).
When Mark Twain visited Kau and Waiohinu in 1866, he visited the sugar plantations and saw a multitude of fruits available. He tried the cherimoya and called the fruit the most delicious fruit known to man and the pride of the islands. Ever since then, interest in the fruit has drastically increased and has been exported to countries around the world to try.

Domestication History
It has been thought that this tree has origin in the highlands of Ecuador & Peru but with recent molecular data, there may be suggestion that there is a meso-American origin. However, now it is grown in subtropical countries with Spain being its largest producer (Morton, 1987). It is also thought to have spread to Chile and Brazil in ancient times as it is currently naturalized in those countries highlands.
It is thought that because the wild and domesticated versions of cherimoya tree are found all over Peru, the domestication history starts in Peru. Both the wild and domesticated cherimoyas have variations in the fruit size and character of the skin (Popenoe 1914). However, unlike guanabana, the other common fruit similar to the cherimoya, there is no archaeological evidence of the cherimoya in Peru, so it is hard to pinpoint exactly where and when domestication of this tree took place ( 2014). There is only speculation that it was domesticated in ancient Peru during the Moche era because of literature and pottery but beyond that there is not much evidence. It is easy to commonly mistake the Cherimoya seeds with the guanabana seeds in archaeology as they’re both black and round. It is known that previously, the cherimoya was unable to be consumed outside of its origin of domestication or cultivation so South America or Spain because it was easily bruised and a soft fruit. However, after new selection aiming to fix those issues, we have a new fruit that is able to be transported longer distances and remove those limitations (National Research Council, 1989).
Because the conditions to grow the cherimoya tree are so specific and the tree itself is very sensitive to its environment, commercial harvesting is proving to be a difficult task. It is only able to be grown on a commercial scale in specific areas of the world, namely Spain and California (National Research Council, 1989). California’s environmental conditions are just right for the cherimoya tree but as discussed in the life history of the tree, the challenge doesn’t lie just in the growing. In order for successful fruit bearing, the farmer must heavily tend to the tree with labor intensive processes of pruning and hand pollination. Therefore, more research and selection needs to be done to successfully harvest the fruit on a larger commercial scale. This is why the fruit itself is quite expensive in non-native areas.

Cultivation Practices
The fruits are very sensitive to climate. If exposed to cold weather before fully mature then they may ripen incorrectly. If they’re exposed to heavy rains or excessive sun then the fruit may crack open. If the humidity is too high, then they have short storage lives (National Research Committee on Technology Innovation, 1989). It is typically grown in Spain, Guatemala, Bolivia, Peru, and other subtropical regions.
To ensure reliable results of bearing fruit, trees are normally grafted or budded on seedling rootstock and done in the spring before the leaves drop (Robert, 2014). The trees themselves prefer higher elevations to 5000 feet but can bear fruits at lower elevations as low as 800 feet (Love, 2012). Pollination is one of the most difficult tasks in cultivating a cherimoya tree. Since the flowers prevent self-pollination with the dichogamous flowering nature and the configuration of the petals, natural pollination is not reliable or conducive. Therefore, hand-pollination is essential in ensuring everything gets pollinated. The pollen would be collected in the evening, stored in a cool, dry place and then reapplied to the mature stigmas in the morning (Robert, 2014).

Nutritional Value and Part of the Plant Consumed
The fruit of the tree or the cherimoya is the part that is consumed. The enticing quality of the fruit is its custardy, creamy texture of the flesh, that give it the title of the custard apple (Arakelyan, 2021).The aromatic flavor is said to be a mix of banana, papaya, and pineapple. It is common to eat the ripe fruit straight out of the tree, cut open, and scooped with a spoon. It also has high nutritional value of around 1.5g of fiber, 21.7g of Calcium, 30.2 mg of Phosphorus, 0.117mg of Thiamine, 0.112g of Riboflavin, and 16.8mg of Ascorbic Acid (Morton, 1987).The cherimoya has micronutrients such as Magnesium, Calcium, and Phosphorus that are in two to four-fold higher in value than other common fruits such as apples. Cherimoya also serves as a great source of Ascorbic Acid (Gentile, 2021).Ascorbic Acid or otherwise known as Vitamin C is a nutrient your body uses to form blood vessels or collagen in bones. It is an essential vitamin to the body’s healing process. Other vitamins such as B6, B2, B1 are all essential in maintaining a healthy immune system and keeping the nervous system functioning. Copper, Potassium, and Magnesium are important minerals help to form blood vessels or are minerals used in upkeeping the body cells.

Health benefits and risks
The seeds should not be eaten since they can contain harmful side effects. The crushed seeds have been used as insecticide. There are also accounts of blindness resulting from the juice of the crushed seeds coming in contact with the eyes. This may be due to the chemical alkaloids found in the seeds such as reticuline, anonaine, lanuginosine, and liriodenine. Human ingestion of the seed can also cause intense photophobia, dilated pupils, vomiting, nausea, dryness in the mouth, flatulence, and other symptoms resembling the effects of a prescription medicine Atropine, used to treat the symptoms of low heart rate or as an antidote for overdose of mushroom poisoning (Morton, 1987). In Mexican culture, the seeds are sometimes pulverized, mixed with grease, and applied on skin for parasitic skin disorders or lice. The skin of the fruit is also used to make a decoction for relieving pneumonia (Morton, 1987). The fruit itself is being studied as possibilities to remedy and offset medical conditions. Cherimoya is high in antioxidants which combat the oxidative stress on the body, lowering risks of cancer and heart disease. Cherimoya’s carotenoid antioxidants such as lutein may also boost eye health as lutein is the main antioxidant in the eyes that is essential in maintaining healthy eyes (Arakelyan 2021).. Also, since cherimoya has high macronutrient content of Potassium and Magnesium, consumption may prevent high blood pressure as those nutrients promote dilation of blood vessels to decrease blood pressure.
Studies suggest Annona cherimoya has anti-cancer properties and the consumption of the fruit can lower blood sugar and cholesterol levels. There has been research done on the leaves of the cherimoya that can show cytotoxicity against microbial species and cell lines of breast, colon, and liver cancer cells. The Annoncaeous acetogenins found in the cherimoya seeds can have a potency of 10,000 times stronger than Adriamycin, a chemotherapeutic drug (Haykal, 2019).
There have also been slight connections with the phytochemicals found in the seed of the cherimoya and the linkage to the West Indian parkinsonian syndrome. This is because the alkaloid extracted from the seed is shown to be neurotoxic to the neurons that release dopamine and that can potentially cause dysfunction and degeneration of those neurons. This effect is then potentially linked to an increased probability of developing West Indian Parkinson disease (Haykal, 2019).

References:
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Richardson, A. C., and P. A. Anderson. (1996). “Hand Pollination Effects on the Set and Development of Cherimoya (Annona Cherimola) Fruit in a Humid Climate.” Scientia Horticulturae (Netherlands).
Schroeder, C. A. (1943). “Hand Pollination Studies on the Cherimoya.” Proceedings. American Society for Horticultural Science, 1943 43:39–41.
Vieth, Robert. (2014). "Cherimoya." University of California Cooperative Extension, Agriculture and Natural Resources, Ventura County, ucanr.edu .
Zonneveld, Maarten van, Xavier Scheldeman, Pilar Escribano, María A. Viruel, Patrick Van Damme, Willman Garcia, César Tapia, José Romero, Manuel Sigueñas, and José I. Hormaza. (2012). “Mapping Genetic Diversity of Cherimoya (Annona Cherimola Mill.): Application of Spatial Analysis for Conservation and Use of Plant Genetic Resources.” PloS One 7(1):e29845. doi: 10.1371/journal.pone.0029845.