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Camu Camu Reproductive Morphology

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Fruit Pulp Nicole Phelan
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Camu Camu: The Super Fruit of the Amazon Basin Holland James Smith Myrciaria dubia (Kunth) McVaugh or camu camu, is a tropical shrub-like tree that is best-known by its highly acidic, nutrient-rich berries. The plant has been extremely important for the Yanomamo and Kayapo Indigenous peoples of the Amazon Basin, who have harvested them for thousands of years. Wild and domesticated populations alike are harvested by guiding canoes through the branches during the wet season. The camu camu plant grows to be about 4–8 meters tall, and both the leaves and berries are harvested for their nutritional and medicinal properties. Like many other species within the Myrtaceae family, camu camu evolved within the Amazon Basin, and grows along the rocky riverbeds and swamps in Brazil, Venezuela, Peru, Colombia, and Bolivia. There are significant gaps in knowledge in the phylogenetic history of the plant, which creates avenues for future research. In terms of phenology, camu camu plants synchronize their growth cycles according to the flood-levels of these watersheds, producing small bisexual white flowers during periods with low water levels. As the water levels rise, the plant transitions into its fruiting stage, where it produces small, reddish-purple fleshy fruits. The camu camu berries are known for their high vitamin C content, and small amounts of various amino acids and fatty acids. The berries of camu camu have many anti-inflammatory and antimicrobial properties, and they are a valuable food source to the birds, mammals, fish, and people living in the Amazon Basin. The growing scientific studies on the plant's medicinal properties have increased demand for the fruit globally. The growth in international markets has led farmers to harvest more camu camu berries, negatively impacting wild populations. By Nicole Phelan and Holland Smith #CamuCamu
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Camu Camu: The Super Fruit of the Amazon Basin Myrciaria dubia (Kunth) McVaugh or camu camu, is a tropical shrub-like tree that is best-known by its highly acidic, nutrient-rich berries. The plant has been extremely important for the Yanomamo and Kayapo Indigenous peoples of the Amazon Basin, who have harvested them for thousands of years. Wild and domesticated populations alike are harvested by guiding canoes through the branches during the wet season. The camu camu plant grows to be about 4–8 meters tall, and both the leaves and berries are harvested for their nutritional and medicinal properties. Like many other species within the Myrtaceae family, camu camu evolved within the Amazon Basin, and grows along the rocky riverbeds and swamps in Brazil, Venezuela, Peru, Colombia, and Bolivia. There are significant gaps in knowledge in the phylogenetic history of the plant, which creates avenues for future research. In terms of phenology, camu camu plants synchronize their growth cycles according to the flood-levels of these watersheds, producing small bisexual white flowers during periods with low water levels. As the water levels rise, the plant transitions into its fruiting stage, where it produces small, reddish-purple fleshy fruits. The camu camu berries are known for their high vitamin C content, and small amounts of various amino acids and fatty acids. The berries of camu camu have many anti-inflammatory and antimicrobial properties, and they are a valuable food source to the birds, mammals, fish, and people living in the Amazon Basin. The growing scientific studies on the plant's medicinal properties have increased demand for the fruit globally. The growth in international markets has led farmers to harvest more camu camu berries, negatively impacting wild populations. By Nicole Phelan and Holland Smith #CamuCamu Nomenclatural History
   Myrciaria dubia (Kunth) McVaugh, commonly known as camu camu, is a shrub-like tree in the Myrtaceae family that is native to the Amazon Basin tropics. Camu camu typically grows along watersheds in Peru, Venezuela, Brazil, Bolivia, and Columbia (Engels & Brinckmann, 2012). While on an exploration across the Americas in 1823, Alexander von Humboldt, Aimè Bonpland, and German botanist Karl S. Kunth collected the type specimen of camu camu in Artures, Venezuela, assigning it the Linnaean name Psidium dubium (Tropicos). Later, the plant was renamed Myrciaria dubia by American botanist Robert McVaugh in 1963 (Tropicos). Other common names in European countries include "rumberry," in English, and "araza de aqua," in Spanish. Shifting from the Eurocentric view, the plant has been an important part of local Indigenous cultures prior to European discovery and identification. In addition to “camu camu,” the plant and its fruit have many other common and indigenous names. Areas in Peru typically use "camo camo" and "camu-camu negro." "Azedinha," "cacari," "miraúba," and "muraúba" are commonly used in Brazil. There are even similarities between the names used in Bolivia ("guapuro blanco") and Venezuela ("algracia," "guayabillo blanco," "guayabito," and "limoncillo") (Engels & Brinckmann, 2012).

Evolutionary History
   Myrciaria dubia is a eudicot in the Myrtaceae family, which includes approximately 121 genera and 5800 species of tropical perennial trees and shrubs that produce fleshy fruits (de Paulo Farias et al., 2020). Plants in the Myrtaceae family are known particularly for their nutrient-dense, antioxidant-rich berries, which typically have a glossy exterior and deep purple mesocarp when ripe (de Paulo Farias et al., 2020). Fruits in the Myrtaceae family are also high in carotenoids, phenolic compounds, and volatile compounds, which contribute to the fruit’s medicinal properties (de Paulo Farias et al., 2020). Zooming in, the Myrciaria genus includes approximately 99 species native to the Amazon Basin, Brazil, Bolivia, Paraguay, Argentina, Central America, and South Florida (Borges et al., 2014). Myrciaria berries are deep-purple to red in color, and both the leaves and the fruits are consumed for their nutritional and medicinal value (Borges et al., 2014). In terms of systematics, the taxa within the Myrtaceae family have been historically understudied, and future research is necessary for forming distinct hypotheses of the Myrtaceae phylogeny and geographic origins of species (Lucas et al., 2005).

Ecology, Life History, and Wild Ancestors
   The phenology of the camu camu plant varies between natural and cultivated populations, based on differences in rainfall and river behavior (Ferreira et al., 2021). In a study comparing wild phenology and cultivated phenology, wild populations exhibited the most synchronized flowering in November and December, whereas cultivated populations had the most synchronized flowering in October (Ferreira et al., 2021). In both populations, flowering occurred during periods with higher sun exposure, lower temperatures, and lower water levels in the river (Ferreira et al., 2021). This indicates that flooding patterns and light availability are critical to influencing the reproductive patterns of camu camu. It is also important to note that the camu camu flowers produced during this period are bisexual and self-compatible. However, because the pistil and stamens do not synchronize their growth, the flowering phenology promotes outcrossing (Castro et al., 2018).
  The camu camu flowers transition into the fully ripened purple-red fruit over the course of 88–102 days (Neves et al., 2015). In the study comparing wild and cultivated phenology, both wild and cultivated populations began producing green fruits when river levels began to rise. This occurred between January and March for wild plants and between December and April for cultivated plants (Ferreira et al., 2021). The wild fruits commonly ripen between January and May, whereas the cultivated fruits ripened between January and April (Ferreira et al., 2021). The camu camu berries are a valuable food source for many of the fish and bird populations near the riverbanks, including the fish species Colossoma macropomum, an important indigenous food source commonly known as "gamitana" (Penn JR, 2006). The fish, bird, and human populations that consume the berries are critical for aiding in the seed dispersal of the plant.
   Camu camu is also incredibly vulnerable to disease, insect infestations, fungal infestations, and other pests within their natural habitats. The most prominent pests for camu camu, which are found in over 50% of cultivated camu camu crops, include Tuthillia cognata, Xylosandrus compactus, Conotrachelus dubiae, and Edessa sp. (Penn JR, 2006). The parasitic plant Moradendron spp. is much less common than insect pests, but farmers consider it a higher threat because it interferes with camu camu growth cycles (Penn JR, 2006). As a result, genetic diversity within camu camu populations is important for maintaining ecosystem stability and population resilience.

Reproductive and Vegetative Morphology
   Myrciaria dubia is a woody shrub that can grow to be 4–8 meters in height, with many branching secondary stems (Castro et al., 2018). The trunk and branches are reddish-brown with a smooth texture, and they can grow up to 15 centimeters in diameter (Black, 2021). During drought periods, the outer layer of the bark naturally peels off (Castro et al., 2018). Camu camu produces simple, broad leaves that are approximately 3–10 centimeters long and 1.5–4.5 centimeters wide (Black, 2021). Depending on the individual plant, the leaves are generally elliptic or lanceolate in shape with one midvein and 18–20 reticulate secondary veins (Castro et al., 2018). The leaf phyllotaxy is opposite, which means that there are two opposite leaf nodes at a single point on the branch (Black, 2021). Despite being a perennial species, camu camu plants shed and grow new leaves based on seasonality. In periods of low rainfall, camu camu experiences the greatest amount of new leaf growth (Ferreira et al., 2021). However, during periods of peak flooding, the camu camu plant slows the production of new leaves and mostly consists of mature leaves (Ferreira et al., 2021).
   Myrciaria dubia is an angiosperm that produces small bisexual white flowers that have 4 rounded petals, that are approximately 3–4 millimeters long and have a ciliated margin (Castro et al., 2018). The camu camu flowers have 4 sepals that are approximately 2 millimeters long, 125 stamens that are approximately 6–10 millimeters long, and 1 pistil that is approximately 10–11 millimeters long (Castro et al., 2018). All of the structures within the camu camu flower are free and unfused. Upon fertilization, the camu camu plants produce small fleshy fruits that are 1–5 millimeters in diameter (Castro et al., 2018). The camu camu peel is light yellow-green during the beginning of its growth, and it becomes redder as the fruit ripens. The fully mature berry has a deep reddish-purple peel, which has a high vitamin C content (Castro et al., 2018). Each individual camu camu berry has between 1–4 seeds that are rich in phenolic compounds including flavonoids, phenolic acids, tannins, stilbenes, and lignans (Langley et al., 2015). When consumed, the seeds offer increased anti-inflammatory compared to other tropical fruits (Langley et al., 2015). The pulp of the fruit is pink in color and highly acidic (Black, 2021).

Historical Accounts
   Myrciaria dubia has been intrinsically tied to the indigenous peoples of the Amazon Basin prior to European colonization for thousands of years. Indigenous peoples have been harvesting camu camu berries, leaves, and timber for medicinal uses. Globally, from the initial colonization of the region to the 1970’s, Myrciaria dubia was harvested for timber. Significant scientific research was conducted on the plant and its medicinal properties starting in the 1970’s, prompting an explosion in international market demand. Since then, the fruit has become a staple export from the countries it natively grows, and it is still used by locals for medicinal and culinary purposes (Engels & Brinckmann, 2012).

Domestication History
  While the domestication history of Myrciaria dubia is largely unknown, there are some wild and domesticated populations of the plants, mixed throughout the water system of the region. Indigenous domestication practices aim to mimic the natural ecology of the plant, and as such are planted along rocky riverbanks. Genetic analysis has revealed that there are no major differences between wild and artificial/domesticated populations, as the local people do not select for specific traits (Castro et al., 2018). It is worth noting that many of the wild patches of Myrciaria dubia are upriver of the domesticated plants, possibly explaining the genetic similarity (Šmíd et al., 2017).

Cultivation Practices
   Myrciaria dubia naturally grows along the rocky banks of rivers and lakes along the Amazon Basin river ecosystem. The plant is harvested annually during the wet season, when the base of the trunks are submerged under several feet of water. The berries and leaves are harvested by guiding canoes through the branches and collecting them by hand. Among the myriad of articles and studies on Myrciaria dubia, there are no mentions of the historical Indigenous groups who cultivated it, but the Yanomamo and Kayapo peoples have been living in the areas it natively grows for thousands of years (World Wide Fund for Nature).

Nutritional Value and Part of the Plant Consumed
   When managed properly, almost the entirety of the Myrciaria dubia plant can be used by humans. Although camu camu berries are sometimes consumed raw, the high acidity makes them difficult to eat alone. In cities and towns within the Amazon Basin region, it is common to consume camu camu berries in sweet drinks or on ice cream (Engels & Brinckmann, 2012). Fresh camu camu berries do not maintain their beneficial compounds and vitamins, so they are commonly processed into a powder to remove the highly-acidic taste for international markets. The camu camu berry is unique in its extremely high vitamin C content, with 100 times more vitamin C compared to an equal quantity of oranges (Castro et al., 2018). The berries also contain small amounts of various beneficial essential amino acids, essential fatty acids, minerals, and vitamins. Other parts of the plant are used medicinally, including the tree bark of camu camu plants which can processed into a poultice and applied to wounds to prevent infection or dried and used in a disinfectant practice (Camu-Camu Berries: History, Gastronomical Uses and Nutrition, 2012). On top of all that, the leaves are also used as an herbal remedy (Engels & Brinckmann, 2012). Studies have shown that the leaves contain various beneficial chemicals such as a-pinene and limonene, which have anticancer properties (Pino & Quijano, 2011).

Human Experience with Camu Camu
   Myrciaria dubia has been deeply intertwined with Amazon Rainforest Indigenous peoples for thousands of years, proving to be a powerful and resilient plant. Currently, Myrciaria dubia have been increasingly overexploited due to its growing international market, causing changes in cultivation practices. Local agroforestry farmers have monetary incentives to harvest more berries causing a shift in the fecundity of new saplings, having a long-term negative effect on population density and the ability to sustainably harvest them. Despite the small-scale agriculture of the plant, studies have still shown a significant decrease in the quantity and efficiency of wild populations (Peters, 2018). Although Myrciaria dubia is helping local people and farmers economically, it is unknown what cost it will have on the surrounding environments and towns when these crops become inaccessible or the growing conditions become unsuitable. While there are efforts to preserve wild populations, future efforts should be made to industrially cultivate camu camu berries and increase regulation on the quantity harvested from wild sites, allowing native populations to passively rebound. Although multiple scientific studies have been conducted on Myrciaria dubia medicinal value, additional research is still required, as its “ beneficial effects include antioxidative and anti-inflammatory activities, antiobesity, hypolipidemic, antihypertensive and anti-diabetic effects, DNA damage and cancer protection effects, and other bioactivities'' (Castro et al., 2018). We have only scratched the surface of the services Myrciaria dubia provides for humans, but in order to sustain those benefits we need to preserve and maintain healthy wild populations, as well as the traditional ecological knowledge tied to camu camu.

References
Black, J. (2021). What is Camu Camu (Myrciaria dubia). Herbs America Inc. https://herbs-america.com/what-is-camu-camu/#fn2-13598

Borges, L. L., Conceição, E. C., & Silveira, D. (2014). Active compounds and medicinal properties of Myrciaria genus. Food Chemistry, 153, 224–233. https://doi.org/https://doi.org/10.1016/j.foodchem.2013.12.064

Castro, J. C., Maddox, J. D., & Imán, S. A. (2018). Camu-camu—Myrciaria dubia (Kunth) McVaugh. In S. Rodrigues, E. de Oliveira Silva, & E. S. de Brito (Eds.), Exotic Fruits (pp. 97–105). Academic Press. https://doi.org/https://doi.org/10.1016/B978-0-12-803138-4.00014-9

de Paulo Farias, D., Neri-Numa, I. A. de Araújo, F. F., & Pastore, G. M. (2020). A critical review of some fruit trees from the Myrtaceae family as promising sources for food applications with functional claims. Food Chemistry, 306, 125630. https://doi.org/https://doi.org/10.1016/j.foodchem.2019.125630

Engels, G., & Brinckmann, J. (2012). Camu-camu Myrciaria dubia Family: Myrtaceae. The Journal of the American Botanical Council, (94), 1–4.

Ferreira, G. A. C., Barnett, A. P. A., & Krug, C. (2021). Phenology and fruit set comparison of camu-camu (Myrciaria dubia) in a natural population and a plantation in the Central Amazon, brazil. Acta Amazonica, 51(2), 91–101. https://doi.org/10.1590/1809-4392202000581

Knoji. (2012). Camu-Camu Berries: History, Gastronomical Uses and Nutrition. https://knoji.com/article/camucamu-berries-history-gastronomical-uses-and-nutrition/ Langley, P. C., Pergolizzi, J. V, Taylor, R., & Ridgway, C. (2015). Antioxidant and Associated Capacities of Camu Camu (Myrciaria dubia): A Systematic Review. The Journal of Alternative and Complementary Medicine, 21(1), 8–14. https://doi.org/10.1089/acm.2014.0130

Lucas, E. J., Belsham, S. R., Lughadha, E. M. N., Orlovich, D. A., Sakuragui, C. M., Chase, M. W., & Wilson, P. G. (2005). Phylogenetic patterns in the fleshy-fruited Myrtaceae – preliminary molecular evidence. Plant Systematics and Evolution, 251(1), 35–51. https://doi.org/10.1007/s00606-004-0164-9

Neves, L. C., da Silva, V. X., Chagas, E. A., Lima, C. G. B., & Roberto, S. R. (2015). Determining the harvest time of camu-camu [Myrciaria dubia (H.B.K.) McVaugh] using measured pre-harvest attributes. Scientia Horticulturae, 186, 15–23. https://doi.org/https://doi.org/10.1016/j.scienta.2015.02.006

Penn JR, J. W. (2006). THE CULTIVATION OF CAMU CAMU (MYRCIARIA DUBIA): A TREE PLANTING PROGRAMME IN THE PERUVIAN AMAZON. Forests, Trees and Livelihoods, 16(1), 85–101. https://doi.org/10.1080/14728028.2006.9752547

Peters, C. M. (2018). Wild Camu-Camu of Peru. Mother Earth Gardener. https://www.motherearthgardener.com/plant-profiles/wild-camu-camu-ze0z1802zphe/ Pino, J. A., & Quijano, C. E.(2008). Volatile Constituents of Camu-camu (Myrciaria dubia (HBK) McVaugh) Leaves. Journal of Essential Oil Research, 20(3), 205–207. https://doi.org/10.1080/10412905.2008.9699991

Šmíd, J., Kalousová, M., Mandák, B., Houška, J., Chládová, A., Pinedo, M., & Lojka, B. (2017). Morphological and genetic diversity of camu-camu [Myrciaria dubia (Kunth) McVaugh] in the Peruvian Amazon. PloS one, 12(6), e0179886. https://doi.org/10.1371/journal.pone.0179886

Tropicos. (n.d.). www.tropicos.org/Specimen/101149789

World Wide Fund for Nature. (n.d.) Amazon People https://wwf.panda.org/discover/knowledge_hub/where_we_work/amazon/about_the_amazon/people_amazon/