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Plants & People: The Intersection of Botany and Human Experience

Carob: From Pod to Pantry

Nomenclatural History

The scientific name Ceratonia siliqua L. is derived from the Greek work keras, meaning “horn”, and the Latin word siliqua, which refers to the shape and fitness of the pod. And, of course, the L. is in reference to Carl Linnaeus. The binomial name was first recorded by Linnaeus in Species Plantarum 2 in May 1753, with a locality of Apulia and Sicily (Italy), Crete (Greece), Cyprus, Syria, and Palestine. The designated lectotype is located at the Linnean Society of London Herbarium (Missouri Botanical Garden, n.d.). The common name for this plant is carob, and has its roots in ancient Mediterranean/Middle Eastern cultures. Carob originates from the Hebrew word kharuv, which means “locust”. This relates to two other common colloquial names for carob: St. John’s bread and locust bean. These two names stem from the biblical references to St. John the Baptist’s consumption of locusts during his travels. The “locusts” he was consuming were likely actually the pods from carob trees, hence the adoption of common names that have to do with the word locust. As the carob tree spread throughout the Mediterranean and other parts of the globe, other names were adopted in the new regions, such as the Arabic kharrūb, Spanish algarrobo or garrofero, Italian carrubo, and many more (Batlle & Tous, 1997, p. 9).

Evolution & Domestication

Evolutionary History
Ceratonia siliqua is a member of the Leguminosae family, the third largest angiosperm family and possibly originating in the late Cretaceous period (110.5-66 mya). (Wojciechowski et al, 2004 ). Within this classification, carob belongs to the subfamily Caesalpinioideae which, along with the other two Leguminosae clades (Mimosoideae and Papilionoideae), is evidenced to have been present since the Eocene (55.8-33.9 mya) (Wojciechowski et al, 2004). C. siliqua is then a member of the Caesalpinioideae subfamily that diverged even further into a smaller group known as the Umtiza Clade, to which C. siliqua belongs. The use of fossil records has allowed researchers to estimate that the Umtiza Clade was likely present as far back as the Late Paleocene, dating it to about 56.5 mya, making it one of the oldest Caesalpinioid clades. (Bruneau et al, 2008).
It was long thought that C. siliqua was the only member of the genus Ceratonia, but in the mid-1900s another species of the Ceratonia genus was discovered: C. oreothauma, which is native to Arabia and Somalia. C. oreothauma has been proposed as the potential wild ancestor of the cultivated C. siliqua due to its smaller, less evolved pollen form. It is distinct from the cultivated species since its flowers are strictly unisexual. (Batlle & Tous, 1997, p. 9, 14).

Domestication History

Ecology & Life History

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Morphology

Vegetative Morphology
Carob is a sclerophyllous evergreen shrub or tree with a thick trunk with rough bark, and sturdy branches that may grow up to approximately 10 m high, though wild trees may grow a few meters taller (Batlle & Tous, 1997, p. 10). Some features of sclerophyllous trees include hard, leathery leaves and short internodal distances, which allow them to adapt to long periods of drought and heat (Alm, 2016). The leaves of the carob tree are 10-20 cm long, alternate, pinnately compound with 4-10 oppositely arranged pairs of leaflets, with or without a terminal leaflet. Each leaflet is 3-7 cm long, ovate or elliptic, dark green on the top with slightly undulate margins. The sclerophyllous leaves have a thick epidermis layer to protect from water loss (Batlle & Tous, 1997, p. 10). Carob trees also have a broad, semi-spherical crown that requires low maintenance and are valued as an ornamental shade tree throughout California, Australia, and elsewhere. However, their strong, extensive root system that allows them to adapt to dry conditions in the wild are considered invasive in landscaping (Batlle & Tous, 1997, p. 28).

Reproductive Morphology
Most carob trees are dioecious with female and male flowers on separate trees, though a small number are hermaphroditic. The prevalence of flower types varies by region. For example, hermaphrodite trees are found more frequently in the Mediterranean Spanish coast than in Southern Spain, though they are still rare with respect to male and female trees. Generally, the flowers are 6-12 mm long, arranged along an inflorescence axis in racemes, with coloration that ranges from green to red. They display radial symmetry with no petals and a disc-shaped calyx consisting of 5 hairy sepals. Flowers typically begin bisexual but one sex is repressed during later development, hence the presence of rudimentary stamens or pistils on females or males, respectively. All three types have a nectarial disc, though females produce the highest volume of nectar (Batlle & Tous, 1997, p. 12).

Female flowers feature a fully developed pistil that is 6-8.5 cm long with 2 stigmatic lobes, 2 carpals, and 5 rudimentary stamens. Male flowers have 5 stamens and a central rudimentary pistil. The filaments may either be of either short or long type, while the anthers may range from yellow to red. Hermaphrodite flowers are a combination of both male and female types with a fully developed pistil and 5 fully developed stamens (Batlle & Tous, 1997, p. 12, 14).

The carob fruit is an indehiscent pod or legume 10-30 cm long, 1.5-3.5 cm wide, and about 1 cm thick; they are compressed, with a straight or curved shape depending on the cultivar. Pods are soft and green but become hard, brown, and leathery upon ripening. Carob pods have two main components: the pulp and the seeds. The pulp consists of an outer, dark and leathery pericarp and an inner, soft mesocarp that separates the seeds. The seeds range from 8-10 mm long, 7-8 mm wide, and 3-5 mm thick. They are composed of a hard external coat, an embryo/germ, and the endosperm (Batlle & Tous, 1997, p. 12, 24).

Cultivation Practices

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Consumption & Nutritional Value

Carob is commonly known and utilized as a cacao substitute to its high sugar content (48-56%) and lack of theobromine and caffeine. Currently, the main use is for gum extraction. However, carob pods are an excellent source of bioactive compounds and contain low levels of fat (0.2-0.6%). To get a better idea of the nutritional value carob offers and its wide variety of uses, it’s convenient to split it into its two components. As mentioned previously, the carob fruit can be split upon crushing the pod into the pulp and the seeds, which approximately make up 90% and 10% of the pod’s weight, respectively (Batlle & Tous, 1997, p. 23).

Pulp
The pulp can be ground into large pieces called kibbles, or ground into powder. Kibbles are typically used for animal feed, carob syrup or liqueur, while the powder is used as a cacao substitute or flour for baked goods or sweets. The pulp can also be roasted whole and eaten as a snack (Batlle & Tous, 1997, p. 26-27).
Carob pulp is high in sugar content (48-56%), but also contains cyclitols, fiber, polyphenols such as tannins, amino acids, minerals, and vitamins. The main cyclitol in the pulp is D-pinitol, which has anti-diabetic effects due to the compound's ability to increase insulin sensitivity and regulate blood sugar levels. The fiber and polyphenols in carob are also known to enhance lipid metabolism, lower total and LDL cholesterol to counter hyperlipidemia, and produce anticancer effects that involve anti-tumor, anti-proliferative, and proapoptotic activity. The main polyphenols are tannins, which have anti-diarrheal effects. Carob pulp is also considered a good source of amino acids by the World Health Organization as it contains all 7 essential amino acids at levels that meet their standards. Carob is also a strong source of calcium, and potassium, though it also contains other nutritionally important minerals such as iron, copper, zinc, or manganese (Goulas et al., 2016). Some prominent vitamins in carob pulp include vitamin E, D, C, niacin (B3), B6, and folic acid (B9), with lower levels of A, B2, and B12 (Papaefstathiou et al., 2018).

Seeds
Upon de-hulling the seeds, they are composed of the embryo, which is used in germ meal, and the endosperm. The endosperm is ground into carob bean gum (CBG) also known as locust bean gum (LBG), which is the most common use of commercially grown carob. LBG is used as a stabilizer, thickener, or a binder and gelling agent in a wide variety of food products such as ice creams, soups, sauces, cheese, pies, canned meats, or pet food (Batlle & Tous, 1997, p. 26-28). Germ meal is high in protein content (50%), while LBG is also used for pharmaceutical purposes due to its inhibition of gastrointestinal diseases like gastroesophageal reflux (Batlle & Tous, 1997, p. 24; Goulas et al., 2016).

Human Experience

Historical Applications
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Future Applications
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References

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1. Lepidium meyenii: The fertility root Maca… maybe you’ve heard of it, maybe you consume it on a daily basis. Maca is the highest elevation crop known today, and is native to the high elevation Andes mountains in Peru, growing in elevations up to 4500 m high! Maca has been a highly valued commodity for many indigenous peoples within the Andes mountains of Peru throughout history and persists to be valued today. Known for its turnip-like shape due to its bulbous hypocotyl, it is highly sought after for its roots. Consuming the roots will result in many notable benefits, such as nutritional benefits, mood improvements, and enhancements in energy, strength… even fertility, all which are backed by numerous studies. These benefits are all due to the numerous nutrients, fibers, bioactive compounds, and natural compounds lepidiline A and B that are contained within Maca. Combined, they all make Maca the nutritional powerhouse that is today. Once this nutritional powerhouse was advertised to the world, it became an immediate hit. The demand for Maca created a market boom and countries started trying to cultivate the plant to meet the demand. One country that cultivated Maca is China. In the high mountains of Yunnan, Maca was successfully cultivated and thus became a common ingredient in Chinese traditional medicine. As such, Maca remains an important dietary source not only in Peru, but as well as worldwide! With more research being done today to further explore what components are in Maca, only time will tell what additional health benefits Maca has to offer us. By Mellanie Gamero and Jacqueline Duong
2. Moringa oleifera: Domestication Histories and Emerging Deviations from Indian Origin The superfood, Moringa oleifera, has a rich history deeply rooted in its native Indian domestication and usage. Also widely known as “The Miracle Tree”, M. oleifera is often recognized by its white flowers and long-seed pods that are greatly sought after. Not only does M. oleifera provide an incomparable food source and multipurpose oil through its pods, but all parts of the plant may be used and consumed. The versatility of this plant is venerated in Indian culture where its consumption and, most importantly, medicinal usage involve all parts of the plant. Due to its spread across other countries, the species and its utilization have evolved to suit individual cultures and their own domestication desires. In the Philippines and North and South Africa, its nutritional value is greatly relied upon to treat malnourishment, purify water, and support community infrastructure while the Indian native medicinal usage is less emphasized and not well known. The Americas, on the other hand, have increasingly toyed with the novel M. oleifera ‘organic-superfood’ craze that tends to remove emphasis from its cultural value and multipurpose utilization to instead fawn over its supposed energy boosts, skin care implementations, and uses in its powder – likely less effective and increasingly more expensive than the original. Though its original purpose grows increasingly colonized, the domestication ventures globally of this superfood continue the legacy of the Moringa oleifera species itself – marked genetic increases in energy production associated with plastic to nucleus gene duplications. As each nation tends to and selects for various aspects of this energy contribution – whether for the nutritional, medicinal, or vanity value – the variation in these M. oleifera may grow towards eventual speciation. By George Vetushko and Lauren Guevara #drumsticktree
3. Salvia hispanica: A Source of Strength Traditionally a key component in Aztec and Mexican culture, chia seeds have been used for centuries for their nutritional, decorative, and medicinal qualities. A common staple in many cultures around the world, these tiny seeds pack a powerful punch and are enjoyed by fitness enthusiasts everywhere. While the seeds themselves are easily recognizable, less well known is their source: a plant known as Salvia hispanica. This flowering, annual herb has a fascinating history and distinctive morphological traits that have evolved over time due to human cultivation. By exploring the myriad ways that this plant has interacted with people over the course of history, we can gain a greater appreciation of how chia seeds have been an important source of strength and inspiration for many. By Joshua Matsuda, Sarah Hoffman, Julia Lindner #Chia
4. 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
5. Breadfruit: The Intrepid Fruit's Journey with Indigenous Peoples and Colonization Breadfruit (Artocarpus altilis (Parkinson) Fosberg) is an intrepid dicot with a rich history and culture surrounding it. Breadfruit trees are a single-trunked tree with a spreading, evergreen canopy. Breadfruit trees are monoecious with both the male and female flowers on the same tree. The male flowers are club shaped, and the female flowers are thousands of small flowers with 2 anthers around a spongy core. The name is derived from the Greek words bread and fruit (artos = bread & karpos = fruit). There are over 2,000 documented names because of its widely spread distribution across the Pacific. A. altilis originated from the South Pacific and was later spread throughout Melanesia, Micronesia, and Polynesia by voyagers. As a result, breadfruit is adapted to a wide variety of climates and conditions in equatorial lowlands. In addition, breadfruit trees thrive in humid tropics and can grow in a variety of soils. Breadfruit became a staple crop because of its nutritional and medicinal value. In the 1700s, breadfruit’s ease of maintenance and nutritional value made it a prime crop, in the eyes of colonizers, to feed African slaves in the Caribbean. Today, breadfruit continues to have an important role in sustainability, fighting malnutrition, and increasing food security. There are research efforts to increase the nutritional value and initiatives to promote breadfruit cultivation in tropical and subtropical regions. For example, The National Tropical Botanical Garden established the Breadfruit Institute in 2003 in hopes of promoting the study of breadfruit for agroforestry, reforestation, global food security and economic development.
6. Smallanthus sonchifolius: The Superfood of the Andes Smallanthus sonchifolius is a plant that in recent years has gained a lot of traction by western and European countries. Its ability to store sugars in the form of carbohydrates has resulted in a very profitable market for yacon cultivators. People have begun looking at this food as a healthier alternative to sugars which has in turn led to its global expansion. A food that was once limited to exist within the Andes has now expanded to other continents. Yacón's human impact has been a part of South American culture for thousands of years, as its is believed to have originated during Incan times. During this time period yacón's consumption was limited to religious events today however, it is a food that is consumed globally. Apart from its sweet tuberous roots, other pars of the plant that are edible are the leaves and the flowers. Together these attributes allow yacón to be known as one of the superfoods from the Andes. By Jessica Manriquez and Amanda Lin #yacón
7. Irvingia Gabonensis: The Untapped Diversity of Dika Plant Irvingia gabonensis is a multipurpose, undomesticated fruit tree native to West and Central Africa. Belonging to the family of Irvingiaceae, this evergreen tree species can grow to a height of 40 meters and attain a diameter of about 120 centimeters. The tree produces mango-like fruits that are well known for their nutritious kernels. Even though the fruit of this plant is also referred to as the African bush mango, the plant has no connection to the actual mango fruit. The dika tree can be found in the dry and wet tropical zones such as Nigeria, Angola, Uganda, Congo, Cameroon, Ghana, Togo, and Benin. It is also known to grow in farmland areas, semi-deciduous forests, and gallery forests. Despite its obscurity on the international level, this species should hardly be considered a minor resource. Almost all parts of the plant have a use from its fruit to its bark. Additionally, it serves as a main source of food and income for indigenous groups in Africa such as the Baka tribe from Central Africa. Due to how prized this plant species is, the dika plant is left untouched by the indigenous people when forests are cleared. Once domesticated, the plant can potentially be used to treat the high levels of malnutrition in Africa. Yet, with increasing local and regional demand for the plant’s kernels, the African bush mango has been facing intense exploitation. Additionally, the disappearance of local wildlife such as elephants are affecting the plant’s dispersal. As a result, the I. gabonensis plant has been classified as a highly endangered species. By Ian Morris and Nikhitha Nair #dika
8. Goji Berries: The Key to Eternal Youth? Goji Berries have become a symbol of the Superfood movement which has continued to gain traction in the US and across the world. Although they have served as a token of this emerging trend, goji berries have been renowned in Asia for their nutritious qualities for many centuries. Its use has been documented for almost 2000 years in traditional Chinese medicine (Gross et al., 2006). The goji industry has skyrocketed during the 21st century as they’ve been recognized for their nutritious qualities. Goji berries have a rich cultural history which has contributed to their claim to fame. In 2003, health writer Earl Mindell distributed a booklet which suggested that goji berries had anti-cancer properties - this drove their entry into the Superfood market. His claims were based on the myth of Li Qing Yuen; the herbalist supposedly consumed goji berries daily which allowed him to live 256 years (Earl Mindell and Rick Handel, 2003). While Mindell’s booklet was full of false exaggerations, there is merit to many of his claims regarding the nutritional benefits of goji berries. Two species are colloquially known as goji berries: Lycium barbarum and Lycium chinense. Their unique vegetative and reproductive morphology makes harvesting the berries labor intensive. Both the goji berry and immature leaves from L. chinense and L. barbarum are eaten by many for their health benefits - while the benefits have been known for centuries, the underlying macromolecular structure that dictates these properties is only being explored in more recent decades. By Ismael Rodriguez & Catherine Nordstrom #Goji
9. Tiger Nuts: Cultivar, Weed, and the Plasticity of Plant-People Relationships Contrary to its name, the tiger nut is the edible tuber of Cyperus esculentus, a sedge that originates from the Mediterranean area. While it was considered a valuable crop in pre-dynastic Egypt where it was even entombed with Egyptian officials, Cyperus esculentus became widely known as an invasive weed during the 19th century. Because of its low yield, it was never considered valuable enough to cultivate as a cash crop. Once again, however, the tiger nut is regaining its popularity but now as a superfood. With its high fiber content and ability to promote gut health, it continues to rise in popularity as one of many superfoods available in the market today. By Phan Tran, Karla Gonzalez
15. Açaí Berry: The Exploitation of Labor in South America The açaí berry is a small fruit, similar in size to a blueberry but deeper purple in color. Native to Eastern Amazonia, the açaí berry has only recently begun to be popularized as a “superfood” in urban communities in the United States within the past 20 years. This superfood claims to be helpful for a variety of major health concerns including, but not limited to, arthritis, weight loss, high cholesterol, and detoxification (NCCIH). Even though research is limited and these claims remain unproven, the popularity of açaí berries still manages to grow exponentially each year. The global market for the açaí berry is expected to exceed $2 billion by 2026 (McCoy). The only proven claim than can be made about the açaí berry is that its rise in popularity only serves to threaten the lives of families, both adults and children, that are living in the surrounding communities in the Amazon who are responsible for harvesting the berries and are, in turn, routinely exploited for cheap, quick labor. More often than not, these açaí pickers, known as peconheiros, are forced to work in critically dangerous conditions and are paid little to nothing. Peconheiros are regularly exposed to serious injuries including bone fractures, accidental knife wounds, and venomous snake and spider bites. With the açaí trees growing upwards of 50 feet tall and only a braided rope being provided to climb, falls are common– and sometimes fatal. Further, peconheiros have no way of advocating for themselves in the event that they do become injured in the forest. No laws or regulations exist that protect the safety and wellness of the peconheiros. by Roberto Vindel, Pauline Le, and Max Kwon #Açaí
17. The Wonder Berry : Phyllanthus emblica Known for being rich in both nutrition and history, the amla berry or as Carl Linneaus named it in 1753, Phyllanthus emblica, can be classified as India’s “Wonder Berry.” Originally believed to come from the semi-arid areas and plains of Northern India, the plant is native to Asia's tropic south-eastern regions such as Vietnam, Laos, and Thailand. Such regions provide conditions like mixed forests and elevations, that the plant cannot live without. However, with the fruit’s beneficial properties being so popular, countries all over the world cultivate the amla berry with India well beyond 100,000 acres of amla development. The amla berry’s influence has even reached as far as the United States where seed distribution first began in the early 1900s. It is said that the Indian god Brahma’s tears germinated the first seeds. In fact, the origin of the fruit’s usage can be traced hundreds of years ago where it was and still used today as a healing remedy. This fruit has been used to cure fevers, joint inflammations, and constipation for hundreds of years. Even recent research has suggested that the antiviral and antibacterial properties of the amla berry may be able to prevent cancer, diabetes, and excessive cholesterol. Despite only being the size of a golf ball, the amla berry is very high in vitamin C, vitamin E, vitamin A, iron, and calcium. In contrast to the size, these berries grow on trees that can reach heights of 12 meters all while surrounded by the beauty of greenish-yellow and pink flowers running across the branchlets. Ultimately, it is safe to say that such a plant is truly a wonder. By Natalie Nartz and Matthew Nguyen #amla
25. From medicine to industry: the multifaceted history of Guaraná (Paullinia cupana Kunth., Sapindaceae) Paullinia Cupana, nature’s caffeine tablet, has a very deep history within native Americans that resided in South America. Known by locals as guarana, “drink of the Lord”, it has a very easily recognizable eye ball like structure, with a large black seed surrounded by bright red fruit. The seed contain an extremely high concentration of caffeine and tanin, four times the amount found in coffee beans. It was commonly used by the Satere Maue people to make beverages and used as medicine for bodily discomfort as far back as over 2000 years ago. Nowadays, despite its popularity, the only country that cultivates guarana is Brazil, with over 15,000 hectares of land growing guarana. It is such an important crop for the people of Brazil, parts of the Amazon was cut down in order to make room to grow guarana. In 2006, the cash value of domestic production of guarana is over 13 billion, and it has only been increasing due to the growing domestic and international demand. Due to it’s high caffeine content within the seed, it is very widely used nowadays to be included in energy drinks. One extremely popular soda called Guarana Antarctica is the biggest competitor of Coca Cola in Brazil. By Tim Sisneros and Edward Teng #Guarana #Paullinia
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