Brassica oleracea explained

Brassica oleracea is a plant species from family Brassicaceae that includes many common cultivars used as vegetables, such as cabbage, broccoli, cauliflower, kale, Brussels sprouts, collard greens, Savoy cabbage, kohlrabi, and gai lan.

Its uncultivated form, wild cabbage, native to coastal southern and western Europe, is a hardy plant with high tolerance for salt and lime. However, its intolerance of competition from other plants typically restricts its natural occurrence to limestone sea cliffs, like the chalk cliffs on both sides of the English Channel.^[1] Wild B. oleracea is a tall biennial plant that forms a stout rosette of large leaves in the first year. The leaves are fleshier and thicker than other Brassica species—an adaptation that helps it store water and nutrients in its difficult growing environment. In its second year, it uses the stored nutrients to produce a flower spike 1to(-) tall with numerous yellow flowers.

A 2021 study suggested that the Eastern Mediterranean Brassica cretica was the origin of domesticated B. oleracea. Genetic analysis of nine wild populations on the French Atlantic coast indicated their common feral origin, deriving from domesticated plants escaped from fields and gardens.^[2]

Taxonomy

Origins

According to the Triangle of U theory, B. oleracea is very closely related to five other species of the genus Brassica.^[3] A 2021 study suggested that Brassica cretica, native to the Eastern Mediterranean, particularly Greece and the Aegean Islands, was the closest living relative of cultivated B. oleracea, thus supporting the view that its cultivation originated in the Eastern Mediterranean region, with later admixture from other Brassica species.^[4]

The cultivars of B. oleracea are grouped by developmental form into several major cultivar groups, of which the Acephala ("non-heading") group remains most like the natural wild cabbage in appearance. For a list of these groups, see the table of cultivars.

Etymology

'Brassica' was Pliny the Elder's name for several cabbage-like plants.^[5]

Its specific epithet oleracea means "vegetable/herbal" in Latin and is a form of (Latin: oleraceus).^{[6] [7]}

Cultivation and uses

B. oleracea has become established as an important human food crop plant, used because of its large food reserves, which are stored over the winter in its leaves. It has been bred into a wide range of cultivars, including cabbage, broccoli, cauliflower, brussels sprouts, collards, and kale, some of which are hardly recognizable as being members of the same genus, let alone species.^[8] The historical genus of Crucifera, meaning "cross-bearing" in reference to the four-petaled flowers, may be the only unifying feature beyond taste.

Researchers believe it has been cultivated for several thousand years, but its history as a domesticated plant is not clear before Greek and Roman times, when it was a well-established garden vegetable. Theophrastus mentions three kinds of (ῤάφανος):^[9] a curly-leaved, a smooth-leaved, and a wild-type.^[10] He reports the antipathy of the cabbage and the grape vine, for the ancients believed cabbages grown near grapes would impart their flavour to the wine.^[11]

History

Through artificial selection for various phenotype traits, the emergence of variations of the plant with drastic differences in looks occurred over centuries. Preference for leaves, terminal bud, lateral bud, stem, and inflorescence resulted in selection of varieties of wild cabbage into the many forms known today.

Impact of preference

• The preference for the eating of the leaves led to the selection of plants with larger leaves being harvested and their seeds planted for the next growth. Around the fifth century BC, the formation of what is now known as kale had developed.^[12]
• Preference led to further artificial selection of kale plants with more tightly bunched leaves, or terminal bud. Somewhere around the first century AD emerged the phenotype variation of B. oleracea known as cabbage.
• Phenotype selection preferences in Germany resulted in a new variation from the kale cultivar. By selecting for fatter stems, the variant plant known as kohlrabi emerged around the first century AD.
• European preference emerged for eating immature buds, selection for inflorescence. Early records in 15th century AD, indicate that early cauliflower and broccoli heading types were found throughout southern Italy and Sicily, although these types may not have been resolved into distinct cultivars until about 100 years later.^{[13] [14] [15]}
• Further selection in Belgium in lateral bud led to Brussels sprouts in the 18th century.

Cultivar groups

According to the Royal Botanic Gardens (Kew Species Profiles)^[16] the species has eight cultivar groups. Each cultivar group has many cultivars, like 'Lacinato' kale or 'Belstar' broccoli.

• Acephala

non-heading cultivars (kale, collards, ornamental cabbage, ornamental kale, flowering kale, tree cabbage).

• Alboglabra

Asian Cuisine cultivars (Chinese kale, Chinese broccoli, gai lan, kai lan).

• Botrytis

cultivars that form compact inflorescences (broccoli, cauliflower, broccoflower, calabrese broccoli, romanesco broccoli).

• Capitata

cabbage and cabbage-like cultivars (cabbage, savoy cabbage, red cabbage).

• Gemmifera

bud-producing cultivars (sprouts, Brussels sprouts)

• Gongylodes

turnip-like cultivars (kohlrabi, knol-kohl)

• Italica

sprouts (purple sprouting broccoli, sprouting broccoli). Edible inflorescences not compacted into a single head.

• Tronchuda

low-growing annuals with spreading leaves (Portuguese cabbage, seakale cabbage).

A 2024 study compares 704 B. oleracea sequences and establishes a phylogenetic tree of cultivars. The authors find large-scale changes in gene expression and gene presence. Some genes are putatively linked to certain traits such as arrested inflorescence (typical of cauliflower and broccoli).^[17]

Cultivar	Image	Cultivar group (Kew)	Name (variety, form)
Wild cabbage		N/A	Brassica oleracea var. oleracea
Cabbage			Brassica oleracea var. capitata f. alba
Savoy cabbage			Brassica oleracea var. capitata f. sabauda
Red cabbage			Brassica oleracea var. capitata f. rubra
Cone cabbage			Brassica oleracea var. capitata f. acuta
Gai lan			Brassica oleracea var. alboglabra
Collard greens			Brassica oleracea var. viridis
Jersey cabbage		Acephala	Brassica oleracea var. longata
Ornamental kale		Acephala	Brassica oleracea var. acephala
Kale			Brassica oleracea var. sabellica
Lacinato kale			Brassica oleracea var. palmifolia
Perpetual kale			Brassica oleracea var. ramosa
Kalette		Hybrid	Brassica oleracea var. viridis x gemmifera
Marrow cabbage		Acephala	Brassica oleracea var. medullosa
Tronchuda kale			Brassica oleracea var. costata
Brussels sprout			Brassica oleracea var. gemmifera
Kohlrabi			Brassica oleracea var. gongylodes
Broccoli			Brassica oleracea var. italica
Cauliflower		Botrytis	Brassica oleracea var. botrytis
Caulini		Botrytis	Brassica oleracea var. botrytis
Romanesco broccoli		Botrytis	Brassica oleracea var. botrytis
Broccoli di Torbole			Brassica oleracea var. botrytis
Broccoflower		Hybrid (within Botrytis)	Brassica oleracea var. botrytis × italica
Broccolini		Hybrid	Brassica oleracea var. italica × alboglabra

Uses

It is rich in essential nutrients, including vitamin C. A diet rich in cruciferous vegetables (e.g., cabbage, broccoli, cauliflower) is linked to a reduced risk of several human cancers.^{[18] [19]}

Human genetics in relation to taste

The TAS2R38 gene encodes a G protein-coupled receptor that functions as a taste receptor, mediated by ligands such as PROP and phenylthiocarbamide that bind to the receptor and initiate signaling that confers various degrees of taste perception. Bitter taste receptors in the TS2R family are also found in gut mucosal and pancreatic cells in humans and rodents. These receptors influence release of hormones involved in appetite regulation, such as peptide YY and glucagon-like peptide-1, and therefore may influence caloric intake and the development of obesity. Thus, bitter taste perception may affect dietary behaviors by influencing both taste preferences and metabolic hormonal regulation.^[20]

Three variants in the TAS2R38 gene – rs713598, rs1726866, and rs10246939 – are in high linkage disequilibrium in most populations and result in amino acid coding changes that lead to a range of bitter taste perception phenotypes. The PAV haplotype is dominant; therefore, individuals with at least one copy of the PAV allele perceive molecules in vegetables that resemble PROP as tasting bitter, and consequently may develop an aversion to bitter vegetables. In contrast, individuals with two AVI haplotypes are bitter non-tasters. PAV and AVI haplotypes are the most common, though other haplotypes exist that confer intermediate bitter taste sensitivity (AAI, AAV, AVV, and PVI). This taste aversion may apply to vegetables in general.^{[20] [21]}

References

• Beckett . Emma . Bitter Taste Genetics – the Relationship to Tasting, Liking, Consumption and Health . Food & Function. 2014 . 5 . 12 . 3040–54 . 10.1039/C4FO00539B . 25286017 . 1959.13/1298053 . free .
• Boxer . E.E. . Garneau . N.L. . Rare haplotypes of the gene TAS2R38 confer bitter taste sensitivity in humans . SpringerPlus. 2015 . 4 . 505. 10.1186/s40064-015-1277-z . 26405625 . 4574037 . free .
• Bufe . B. . The Molecular Basis of Individual Differences in Phenylthiocarbamide and Propylthiouracil Bitterness Perception . Current Biology. 2005 . 15 . 4 . 322–7 . 10.1016/j.cub.2005.01.047 . 15723792 . 1400547 . 2005CBio...15..322B .
• Calancie . Larissa . etal . TAS2R38 Predisposition to Bitter Taste Associated with Differential Changes in Vegetable Intake in Response to a Community-Based Dietary Intervention . . 2018 . 8 . 6 . 2107–2119. 10.1534/g3.118.300547 . 29686110 . 5982837 .
• Duffy . V.B. . etal . Vegetable Intake in College-Aged Adults Is Explained by Oral Sensory Phenotypes and TAS2R38 Genotype . Chemosensory Perception. 2010 . 3 . 3–4 . 137–148 . 10.1007/s12078-010-9079-8 . 21157576 . 3000691 .
• Wieczorek . Martyna. . 28896102 . etal . Bitter Taste of Brassica Vegetables: The Role of Genetic Factors, Receptors, Isothiocyanates, Glucosinolates, and Flavor Context. . . 2018 . 58 . 18 . 3130–3140 . 10.1080/10408398.2017.1353478 . 28718657 .
• Rozengurt . E. . Taste Receptors in the Gastrointestinal Tract. I. Bitter taste receptors and α-gustducin in the mammalian gut . . 7 . 171–177.
• Roura . Eugeni . etal . Variability in Human Bitter Taste Sensitivity to Chemically Diverse Compounds Can Be Accounted for by Differential TAS2R Activation. . Chemical Senses. 21 May 2015 . 40 . 6 . 427–35 . 10.1093/chemse/bjv024 . 25999325 . free .
• Risso . David . etal . Erratum: Corrigendum: Global Diversity in the TAS2R38 Bitter Taste Receptor: Revisiting a Classic Evolutionary PROPosal. . . 27 June 2016 . 6 . 28406 . 10.1038/srep28406 . 27346370 . 4921822 .
• Mennella . Vito . 6501199 . etal . Functionally Distinct Kinesin-13 Family Members Cooperate to Regulate Microtubule Dynamics during Interphase. . Nature Cell Biology. 20 February 2005 . 7 . 3 . 235–45 . 10.1038/ncb1222 . 15723056 .
• Kim . U.K. . etal . Genetics of individual differences in bitter taste perception: lessons from the PTC gene . Clinical Genetics. 2005 . 67 . 4 . 275–80 . 10.1111/j.1399-0004.2004.00361.x . 15733260 . 1639438 .
• Kim . U.K. . 8801457 . etal . Genetics of Human Taste Perception . Journal of Dental Research. 2004 . 83 . 6 . 448–53 . 10.1177/154405910408300603 . 15153450 .
• Khataan . Nora H. . 45496712 . etal . TAS2R38 Genotypes and Phenylthiocarbamide Bitter Taste Perception in a Population of Young Adults. . Journal of Nutrigenetics and Nutrigenomics. 2009 . 2 . 4–5 . 251–6 . 10.1159/000297217 . 20484932 . free .
• Genick . U.K. . etal . Sensitivity of genome-wide-association signals to phenotyping strategy: the PROP-TAS2R38 taste association as a benchmark . PLOS ONE. 2011 . 6. 11 . e27745 . 10.1371/journal.pone.0027745 . 22132133 . 3223210 . 2011PLoSO...627745G . free .

External links

• PROTAbase on Brassica oleracea (Brussels sprouts)
• PROTAbase on Brassica oleracea (cauliflower and broccoli)
• Video Overview of Brassica oleracea

from Untamed Science

Notes and References

1. Snogerup . Sven . Gustafsson . Mats . Bothmer . Roland Von . 1990-01-01 . Brassica sect. Brassica (Brassicaceae) I. Taxonomy and Variation . 3996645 . . 19 . 2 . 271–365.
2. 10.1007/s10722-020-00945-0. Survey and genetic diversity of wild Brassica oleracea L. Germplasm on the Atlantic coast of France. 2020. Maggioni. Lorenzo. von Bothmer. Roland. Poulsen. Gert. Härnström Aloisi. Karolina. 218772995. Genetic Resources and Crop Evolution. 67. 7. 1853–1866. 10568/121870. free.
3. Book: Dixon. G.R.. 2007 . Vegetable brassicas and related crucifers. 978-0-85199-395-9. CABI. Wallingford.
4. Mabry . Makenzie E . Turner-Hissong . Sarah D . Gallagher . Evan Y . McAlvay . Alex C . An . Hong . Edger . Patrick P . Moore . Jonathan D . Pink . David A C . Teakle . Graham R . Stevens . Chris J . Barker . Guy . Labate . Joanne . Fuller . Dorian Q . Allaby . Robin G . Beissinger . Timothy . Decker . Jared E . Gore . Michael A . Pires . J Chris . 2021 . The Evolutionary History of Wild, Domesticated, and Feral Brassica oleracea (Brassicaceae) . . 38 . 10 . 4419–4434 . 10.1093/molbev/msab183 . 34157722 . 8476135 . amp . free .
5. Gledhill, David (2008). "The Names of Plants". Cambridge University Press. (hardback), (paperback). pp 76
6. Book: Parker, Peter . A Little Book of Latin for Gardeners . 2018 . . 978-1-4087-0615-2 . 328 . Latin: oleraceus, holeraceus = relating to vegetables or kitchen garden.
7. Book: Whitney, William Dwight . The Century Dictionary and Cyclopedia . . 1899 . 2856 . Latin: holeraceus, Latin: oleraceus, herb-like, Latin: holus, Latin: olus (Latin: oler-), herbs, vegetables.
8. Stansell . Zachary . Hyma . Katie . Fresnedo-Ramírez . Jonathan . Sun . Qi . Mitchell . Sharon . Björkman . Thomas . Hua . Jian . 2018-07-01 . Genotyping-by-sequencing of Brassica oleracea vegetables reveals unique phylogenetic patterns, population structure and domestication footprints . . en . 5 . 1 . 38 . 10.1038/s41438-018-0040-3 . 2052-7276 . 6026498 . 29977574 . 2018HorR....5...38S . 49552482.
9. Compare Theophrastus; raphanis (ραφανίς), "radish", also a Brassica.
10. Book: Zohary . Daniel . Domestication of Plants in the Old World: The Origin and Spread of Domesticated Plants in Southwest Asia, Europe, and the Mediterranean Basin . Hopf . Maria . Weiss . Ehud . Oxford University Press (OUP)) . 2012 . 978-0199549061 . . 199 . en.
11. Theophrastus, Enquiry into Plants, IV.6.16; Deipnosophistae, I, noting the effects of cabbages on wine and wine-drinkers, also quotes Apollodorus of Carystus: "If they think that our calling it a, while you foreigners call it a krambê, makes any difference to us women!" (on-line English text).
12. Web site: Vegetables - University of Saskatchewan. agbio.usask.ca. 2016-04-07. https://web.archive.org/web/20160329102224/http://agbio.usask.ca/community-outreach/gardenline/vegetables.php. 2016-03-29. dead.
13. Maggioni. Lorenzo. von Bothmer. Roland. Poulsen. Gert. Branca. Ferdinando. 2010-06-01. Origin and Domestication of Cole Crops (Brassica oleracea L.): Linguistic and Literary Considerations1. Economic Botany. en. 64. 2. 109–123. 10.1007/s12231-010-9115-2. 2771884. 1874-9364. 10568/121874. free.
14. Web site: Maggioni. Lorenzo. June 2015. Domestication of Brassica oleracea L.. 2020-11-29. pub.epsilon.slu.se. sv.
15. Stansell. Zachary. Björkman. Thomas. 2020-10-01. From landrace to modern hybrid broccoli: the genomic and morphological domestication syndrome within a diverse B. oleracea collection. Horticulture Research. en. 7. 1. 159. 10.1038/s41438-020-00375-0. 33082966. 7528014. 2020HorR....7..159S . 2052-7276.
16. Web site: Brassica oleracea (wild cabbage) . kew.org . Royal Botanic Gardens. Accessed March 23, 2023  - see "Descriptions" subsection "According to Kew Species Profiles"
17. Li . Xing . Wang . Yong . Cai . Chengcheng . Ji . Jialei . Han . Fengqing . Zhang . Lei . Chen . Shumin . Zhang . Lingkui . Yang . Yinqing . Tang . Qi . Bucher . Johan . Wang . Xuelin . Yang . Limei . Zhuang . Mu . Zhang . Kang . Lv . Honghao . Bonnema . Guusje . Zhang . Yangyong . Cheng . Feng . Large-scale gene expression alterations introduced by structural variation drive morphotype diversification in Brassica oleracea . Nature Genetics . 13 February 2024 . 56 . 3 . 517–529 . 10.1038/s41588-024-01655-4 . free. 10937405 .
18. Verhoeven . D. T. . Goldbohm . R. A. . van Poppel . G. . Verhagen . H. . van den Brandt . P. A. . 1996-09-01 . Epidemiological studies on brassica vegetables and cancer risk . . 5 . 9 . 733–748 . 1055-9965 . 8877066.
19. Higdon . Jane V. . Delage . Barbara . Williams . David E. . Dashwood . Roderick H. . 2007-03-01 . Cruciferous vegetables and human cancer risk: epidemiologic evidence and mechanistic basis . . 55 . 3 . 224–236 . 10.1016/j.phrs.2007.01.009 . 1043-6618 . 2737735 . 17317210.
20. Calancie . Larissa . Keyserling . Thomas C. . Smith-Taillie . Lindsey . Robasky . Kimberly . Patterson . Cam . Ammerman . Alice S. . Schisler . Jonathan C. . TAS2R38 predisposition to bitter taste associated with differential changes in vegetable intake in response to a community-based dietary intervention . . 2018 . 8 . 6 . 2107–2119 . 10.1534/g3.118.300547. 29686110 . 5982837 . Text was copied from the preprint version, which is available under a Creative Commons Attribution 4.0 International License.
21. Behrens . Maik . Gunn . Howard . Ramos . Purita . Genetic, Functional, and Phenotypic Diversity in TAS2R38-Mediated Bitter Taste Perception . . 2013 . 38. 6 . 475–84 . 10.1093/chemse/bjt016 . 23632915 . free .