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Reproductive Morphology of Lupinus mutabilis Sweet

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Inflorescent arrangement of flowers Sarah Simpson
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1. Reproductive Morphology of Lupinus mutabilis Sweet
Petals and flowering Sarah Simpson
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1. Reproductive Morphology of Lupinus mutabilis Sweet
L. mutabilis flowers from the front and side Sarah Simpson
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1. Reproductive Morphology of Lupinus mutabilis Sweet
Seed pod and seed produced. Sarah Simpson
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1. Reproductive Morphology of Lupinus mutabilis Sweet

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Tarwi: A forgotten gift from the Andes Sarah Simpson Tarwi is an Andean lupine that has been cultivated by those native to the Andes for at least 1500 years. The scientific name of this lupine is Lupinus mutabilis, Lupinus meaning “wolfish,” and mutabilis meaning “variable.” It can survive in high elevations ranging from 1650 to 3300 meters. This plant is a dependable crop that has been used by the pre-Inca and Inca people in crop cycling with other native Andean plants such as quinoa and potatoes. Documentation of the use of tarwi has been recorded archeologically in Naza culture tombs and Tiahuanaco ceramics. There are also records of high cultivation of tarwi through Spanish taxation records that recorded 10,000 hectares cultivated. The use of tarwi decreased following Spanish colonization when Old World crops were introduced to the area; by 1980, only 5,200 hectares of tarwi were cultivated in Peru. Tarwi has colorful purple flowers and palmate leaves, making the plant strikingly beautiful. The tarwi plant can self-pollinate and does not rely on one specific pollinator making it less vulnerable to fluctuations in pollinator populations. In addition to being dependable in pollination, tarwi can be planted in poor soil because it can fix nitrogen in the soil. The tarwi plant contains alkaloids, making it less susceptible to pathogens and pests. The seeds of this plant are consumed, but the seeds need to be soaked in water for two to four days to remove the water-soluble alkaloids that make the seeds bitter. As an Andean native plant, Lupinus mutabilis has various properties and uses that have enhanced the diets and lives of Peruvians and other South Americans. By Lysol Patino and Sarah Simpson #tarwi
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Tarwi: A forgotten gift from the Andes Tarwi is an Andean lupine that has been cultivated by those native to the Andes for at least 1500 years. The scientific name of this lupine is Lupinus mutabilis, Lupinus meaning “wolfish,” and mutabilis meaning “variable.” It can survive in high elevations ranging from 1650 to 3300 meters. This plant is a dependable crop that has been used by the pre-Inca and Inca people in crop cycling with other native Andean plants such as quinoa and potatoes. Documentation of the use of tarwi has been recorded archeologically in Naza culture tombs and Tiahuanaco ceramics. There are also records of high cultivation of tarwi through Spanish taxation records that recorded 10,000 hectares cultivated. The use of tarwi decreased following Spanish colonization when Old World crops were introduced to the area; by 1980, only 5,200 hectares of tarwi were cultivated in Peru. Tarwi has colorful purple flowers and palmate leaves, making the plant strikingly beautiful. The tarwi plant can self-pollinate and does not rely on one specific pollinator making it less vulnerable to fluctuations in pollinator populations. In addition to being dependable in pollination, tarwi can be planted in poor soil because it can fix nitrogen in the soil. The tarwi plant contains alkaloids, making it less susceptible to pathogens and pests. The seeds of this plant are consumed, but the seeds need to be soaked in water for two to four days to remove the water-soluble alkaloids that make the seeds bitter. As an Andean native plant, Lupinus mutabilis has various properties and uses that have enhanced the diets and lives of Peruvians and other South Americans. By Lysol Patino and Sarah Simpson #tarwi
Nomenclature and Naming
The linnean binomial nomenclature of this plant is Lupinus mutabilis. Tarwi is in the genus Lupinus and the species is mutabilis. Lupinus is derived from “wolfish” and this refers to the highly organized structure of wolf packs. The term mutabilis is Latin for, mutable, variable, or liable to change. This name could have been chosen because of the variation seen in the colors and seeds that the L. mutabilis plant produces. Some common names for L. mutabilis include: tarwi, chocho, pearl lupin, and Andean lupin. Some of these common names are derived from indigenous naming; called tarwi by the Quechua people and tauri by the Aymara people.
Evolutionary History
L. mutabilis is hypothesized to descend from the wild ancestral parent, L. piurensis. This is hypothesized due to the morphological similarities between L. mutabilis and L. piurensis. Similar morphological traits include glaucous stems, palmate leaf structures, and large colorful flowers. However, a notable difference between the two species is that L. mutabilis has indehiscent pods, large pale seeds, and a higher seed germination rate. It is estimated that L. mutabilis split from its progenitor L. piurensis at around 2600 BP, BP refers to “before present,” also notated as 650 BC. This crop is hypothesized to have been domesticated in the Cajamarca region in Northern Peru. The first archeological evidence of L. mutabilis was found in the Mantaro Valley in Central Peru and dated back to 1800 BP. There has been recent evolutionary history within the L. mutabilis species with the cultivation of new varieties with low alkaloid content. Alkaloids make the taste of the seeds bitter, and seeds would have to be thoroughly washed to remove the flavor. So this led to an interest in breeding a variety of L. mutabilis with low alkaloid content, called L. mutabilis Sweet.

This phylogenetic tree compares the similarities between different lupine species found in the “New World.” This phylogeny documents the appearance of important traits such as digitated leaves and if it is an annual, survive one year, or perennial, survive two years, species. The phylogeny indicates that L. mutabilis has traits of digitated leaves, leaves in the shape of a palm with ‘digits,’ and is an annual species that sprout from seed and dies in one yearly cycle. L. mutabilis shares the trait of digitates leaves with many other Western New World lupine species.
Ecology and Life History
This crop is in the Fabaceae legume family and is an annual crop that dies off after producing bean pods. It is believed to have been domesticated in northern Peru due to its cultivation history that is heavily dependent on the high altitude of the Andes. It usually grows at higher altitudes ranging from 800 to 3000 meters. Although mildly resistant to drought and frosts, prolonged seasons with both will kill the crop.

The growing period lasts anywhere from 150-360 days, and the exact time length depends on the environment. This includes climate, altitude, and how these interact with the genetics of the plant. Development is much like other lupine species, including emergence from the seed, first leaf development, flower development, and pod/seed development. Tawri seeds have a short germination period, producing shoots within days. The flowers develop in mounds, and these mounds are actually flower clusters called a raceme. Raceme are separate flowers attached almost equidistantly by stalks connected to a central stem. A young tarwi plant will have raceme that are just developing, or with only flowers at the base of the stalk fully open, with unopened buds towards the top of the cluster. A mature tarwi plant will have a fully blooming raceme and developing seed pods. It also grows short, branched taproots that interact with rhizobia as the plant ages. Rhizobia are bacteria that exist in the soil that develop symbiotic relationships with legumes by establishing themselves inside the root nodules of the plants. They fix nitrogen that becomes available to the plant and surrounding plants once inside these root nodules, something these plants and bacteria cannot do independently otherwise. Successful inoculation with this bacteria is associated with higher yields. It can enrich the soil with nitrogen that would benefit not only L. mutabilis but other crops grown in rotations with it.
Vegetative and Reproductive Morphology
L. mutabilis has indeterminate growth patterns, meaning that, unlike other plants that stop growing in maturity, tarwi continues growing in height/width. It is also a plant with sympodial branching. It can be a very tall plant, with heights ranging from 0.5-2.8 meters. The leaves are typical and have a palmate structure. One leaf can be divided into 5-12 leaflets, which are lanceolate or oval in shape. A mature tarwi plant’s flowers are blue and purple, with small white banners and a touch of yellow-orange at the center. They grow in long mounds that extend above the rest of the plant. The plant develops seed pods anywhere from 5-10 cm in length. The seeds are about 5 mm in diameter and range from white to black, with mottled varieties of white with brown/black.

Tarwi flowers are arranged in an inflorescence structure that is a highly ordered cluster of flowers along the stem. These flowers come in various colors ranging between white and purple, but tarwi is mainly known for its purple color, which can be used for dyes. These flowers also have yellow banners. The morphology of these flowers includes five petals and corollas that reach between one to two centimeters in length. These flowers are predominately self-pollinating, but they can be cross-pollinated. After pollination, the plant produces bean pods with large seeds that are indehiscent and do not break apart. Each seedpod typically contains four to five seeds. The seeds produced by the L. mutabilis crop contain alkaloids which make them bitter, sometimes toxic, and unfavorable to pathogens and predators.
Historical Accounts
The history of Lupinus mutabilis is less documented than other Lupine species found in the Old World. Although the history of L. mutabilis needs more research, there is still archeological evidence documenting its importance. Tarwi has been cultivated in the Andes for at least 1500 years. There have been archeological findings of Lupinis scattered around Peru. Some evidence of tarwi has been found in Naza culture tombs and Tiahuanaco ceramics. The first finding of L. mutabilis was in Montaro Valley in central Peru and dated back to 1800 BP (Before Present). There have also been archeological findings of a ceremonial vessel from the pre-Inca period that has drawings of tarwi on it. Other than archeological findings, there is recorded evidence of the cultivation of these Lupines through Spanish tax records following the colonization of the New World. This Spanish tax record suggests that tarwi was grown at a large scale, approximated to be 10,000 hectares compared to the modern cultivation of 5,200 hectares in the 1980s.
Nutritional Value and Consumption
The seeds of the tarwi plant are consumed in different dishes. Tarwi contains high protein (46–48%) and oil (16–23%) contents and low amounts of starch. They are comparable to soybeans in their protein content, making them an ideal replacement plant for animal consumption. For these seeds to be edible, they must be soaked in water for two to four days to remove the water-soluble alkaloids in the beans. Otherwise, the beans will be bitter and unpleasant to eat. The beans are then ready to be consumed after being soaked. The texture of the beans is firm and smooth with a nutty taste. These beans have been used in dishes such as ceviche, which is served cold, salads, and soups. These seeds can then be pressed to make an oil that has health benefits. The oil in tarwi seeds contains Oleic and Linoleic acid. Oleic acid is an omega-9 fatty acid that lowers cholesterol and decreases the risk of heart disease. Linoleic acid has similar properties in that it also reduces the risk of heart disease. These acids in the tarwi oil make it a beneficial oil to consume. In addition to consuming the oil, tarwi has also been sold as flour that can be used in making baked goods. Other than consuming the tarwi seeds, the flowers of this plant have also been used to create dyes.
Domestication and Cultivation
The history of South American lupines is interesting due to its complexity and lack of clarity until recently. This is due to the lack of research on the lupine species of the Andes, which have not benefited from a deep understanding of their domestication. All domesticated lupines share similar characteristics that make them favorable for agriculture, including nitrogen fixation and edible parts of the plant. However, L. mutabilis specifically has been relatively hidden, leaving the rest of its history prior to and during domestication to be a mystery. In a recent study by Atchinson et al, researchers collaborated to uncover more about the domestication history of tarwi. The results showed to support that L. mutabilis forms a monophyletic group that is within the clade of L. piurensis, making L. piurensis the most likely living descendant of the original parent plant of tarwi. Additionally, archeological evidence shows that the first tarwi lupine seeds found that is tarwi as we know it today, and not just another wild lupine, is found in a site in Mantaro Valley. This dates back to 1800 BP, and the researchers, therefore, make the estimate that tarwi was domesticated sometime between 1800 BP and 3450 BP by the pre-Incan people of the Andes.
Where, how, and by whom are the plants grown?
Historically, tarwi cultivation has been done only in high elevation areas ranging from 1650 and 3300 meters, such as the Andean highlands. For most of its history, L. mutabilis has been solely cultivated by Andean civilizations, with archeological evidence that it was cultivated by pre-Incan and Incan peoples. Today, it is still Andean growers that grow most of the tarwi grown in the world. Laborers are usually locals that know the crop and live in close proximity to the fields. While there have been recent investments in establishing tarwi cultivation in Europe, these developments are fairly recent and have not yet boomed like other Andean staple crops like quinoa and potatoes. L. mutabilis can be cultivated in rotation with other crops such as potatoes and cereals, as well as other plants that would benefit from its nitrogen fixation abilities. Harvest usually occurs 150-300 days after germination, with the seed pods being almost dry. The yield per hectare is about 500 to 1000 kg under good conditions.
Human Experience and Future Uses
This crop has been vital to those native to the Andes. The tarwi plant was cultivated extensively in the past due to its high protein and oil content. The tarwi plant flourished in poor soil due to its ability to fix nitrogen in the soil, and it was often used in crop cycling with other Andean plants such as quinoa and potatoes. These factors make tarwi a dependable plant. While tarwi is a crop that many have forgotten, people are now actively looking to cultivate it more. There have been community projects centered around the rural cultivation of tarwi in Peru, where the income from these new markets goes back to the Quechua community. The project is centered around giving Quechua women the skills to cultivate tarwi. This project strengthens women's role in their families as they receive income from this cultivation and food that they can bring back to their families. This project aims to help the Quechua people in the province of Huayalas fight against poverty and malnutrition. Outside of South America, tarwi is being recognized as a nutritional crop that can be a replacement for meat. International use could help to shift away from relying on animal consumption and decrease the use of soybeans as a more economical option.
References
Aïnouche, A., Bayer, R.J., & Misset, M.T. (2004). Molecular phylogeny, diversification and character evolution in Lupinus (Fabaceae) with special attention to Mediterranean and African lupines. Plant Systematics and Evolution, 246, 211-222. https://doi.org/10.1007/s00606-004-0149-8

Atchison, G., Nevado, B., Eastwood, R., Contreras-Ortiz, N., Reynel-Rodríguez, C., Madriñán, S., Filatov, D., & Hughes, C. (2016). Lost crops of the Incas: Origins of domestication of the Andean pulse crop tarwi, Lupinus mutabilis. American Journal of Botany, 103. https://doi.org/10.3732/ajb.1600171

Carvajal-Larenas, F. E., Linnemann, A. R., Nout, M. J. R., Koziol, M., & van Boekel, M. A. J. S. (2016). Lupinus mutabilis: Composition, Uses, Toxicology, and Debittering. Critical Reviews in Food Science and Nutrition, 56(9), 1454–1487. https://doi.org/10.1080/10408398.2013.772089

Chalampuente-Flores, D., Bastidas, C. T., & Sørensen, M. (n.d.). The Andean Lupine-‘El Chocho’ or ‘Tarwi’ (Lupinus Mutabilis Sweet). 6.

Cowling, W., Buirchell, B., & Tapia, M. (1998). Lupin. Lupinus spp. Promoting the conservation and use of underutilized and neglected crops. 23.

Croyle, L. (2019, October 23). Foods of Peru: What is Tarwi and How to Eat it. Traveling and Living in Peru. https://www.livinginperu.com/foods-of-peru-what-is-tarwi-and-how-to-eat-it/

Dtarazona. (2009). Español: Ceviche de chochos. Huari, Ancash, Perú. Own work. https://commons.wikimedia.org/w/index.php?curid=8023297

Gulisano, A., Alves, S., Martins, J. N., & Trindade, L. M. (2019). Genetics and Breeding of Lupinus mutabilis: An Emerging Protein Crop. Frontiers in Plant Science, 10. https://www.frontiersin.org/article/10.3389/fpls.2019.01385

Hondelmann, W. The lupin — ancient and modern crop plant. Theoret. Appl. Genetics 68, 1–9 (1984). https://doi.org/10.1007/BF00252301

How tarwi is improving Andean women’s living conditions. (2015, September 6). LifeGate. https://www.lifegate.com/tarwi-andean-women

Klowo_645_fullsize.jpg (841×1200). (n.d.). Retrieved May 26, 2022, from https://storage.googleapis.com/powop-assets/lowo/KLOWO_645_fullsize.jpg

Tarwi (Lupinus mutabilis) – Cultivariable. (n.d.). Retrieved May 26, 2022, from https://www.cultivariable.com/instructions/other-vegetables/how-to-grow-tarwi/

Tarwi_(Lupinus_mutabilis),_cultivated_near_Puno,_Peru_(February_2015).JPG (3056×4592). (n.d.). Retrieved May 26, 2022, from https://upload.wikimedia.org/wikipedia/commons/0/08/Tarwi_%28Lupinus_mutabilis%29%2C_cultivated_near_
Puno%2C_Peru_%28February_2015%29.JPG

This page has annotations:

This page has tags:

This page is referenced by:

Nomenclature and Naming

​​​​​Evolutionary History

Ecology and Life History

Vegetative and Reproductive Morphology

Historical Accounts

Nutritional Value and Consumption

Domestication and Cultivation

Where, how, and by whom are the plants grown?

Human Experience and Future Uses

References

Evolutionary History