Acanthaceae
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Acanthaceae is a
family Family (from la, familia) is a group of people related either by consanguinity (by recognized birth) or affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its members and of society. Idea ...
(the acanthus family) of
dicotyledon The dicotyledons, also known as dicots (or, more rarely, dicotyls), are one of the two groups into which all the flowering plants (angiosperms) were formerly divided. The name refers to one of the typical characteristics of the group: namely, t ...
ous
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants t ...
s containing almost 250 genera and about 2500 species. Most are
tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
herbs In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicina ...
,
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from tree ...
s, or twining vines; some are
epiphytes An epiphyte is an organism that grows on the surface of a plant and derives its moisture and nutrients from the air, rain, water (in marine environments) or from debris accumulating around it. The plants on which epiphytes grow are called phoroph ...
. Only a few species are distributed in
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
regions. The four main centres of distribution are
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Gui ...
and
Malaysia Malaysia ( ; ) is a country in Southeast Asia. The federal constitutional monarchy consists of thirteen states and three federal territories, separated by the South China Sea into two regions: Peninsular Malaysia and Borneo's East Mal ...
,
Africa Africa is the world's second-largest and second-most populous continent, after Asia in both cases. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 6% of Earth's total surface area ...
,
Brazil Brazil ( pt, Brasil; ), officially the Federative Republic of Brazil (Portuguese: ), is the largest country in both South America and Latin America. At and with over 217 million people, Brazil is the world's fifth-largest country by area ...
, and
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
. Representatives of the family can be found in nearly every
habitat In ecology, the term habitat summarises the array of resources, physical and biotic factors that are present in an area, such as to support the survival and reproduction of a particular species. A species habitat can be seen as the physical ...
, including dense or open
forest A forest is an area of land dominated by trees. Hundreds of definitions of forest are used throughout the world, incorporating factors such as tree density, tree height, land use, legal standing, and ecological function. The United Nations' ...
s,
scrubland Shrubland, scrubland, scrub, brush, or bush is a plant community characterized by vegetation dominated by shrubs, often also including grasses, herbs, and geophytes. Shrubland may either occur naturally or be the result of human activity. It ...
s, wet fields and valleys, sea
coast The coast, also known as the coastline or seashore, is defined as the area where land meets the ocean, or as a line that forms the boundary between the land and the coastline. The Earth has around of coastline. Coasts are important zones in n ...
and marine areas,
swamp A swamp is a forested wetland.Keddy, P.A. 2010. Wetland Ecology: Principles and Conservation (2nd edition). Cambridge University Press, Cambridge, UK. 497 p. Swamps are considered to be transition zones because both land and water play a role in ...
s, and
mangrove A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in severa ...
forests.


Description

Plants in this family have simple, opposite,
decussate Decussation is used in biological contexts to describe a crossing (due to the shape of the Roman numeral for ten, an uppercase 'X' (), ). In Latin anatomical terms, the form is used, e.g. . Similarly, the anatomical term chiasma is named af ...
d
leaves A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, st ...
with entire (or sometimes toothed, lobed, or spiny) margins, and without stipules. The leaves may contain
cystolith Cystolith (Gr. "cavity" and "stone") is a botanical term for outgrowths of the epidermal cell wall, usually of calcium carbonate, formed in a cellulose matrix in special cells called lithocysts, generally in the leaf of plants. Cystoliths are pr ...
s, calcium carbonate concretions, seen as streaks on the surface. The
flower A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants (plants of the division Angiospermae). The biological function of a flower is to facilitate reproduction, usually by providing a mechanis ...
s are perfect,
zygomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
to nearly
actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
, and arranged in an
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are forme ...
that is either a spike,
raceme A raceme ( or ) or racemoid is an unbranched, indeterminate type of inflorescence bearing flowers having short floral stalks along the shoots that bear the flowers. The oldest flowers grow close to the base and new flowers are produced as the s ...
, or cyme. Typically, a colorful
bract In botany, a bract is a modified or specialized leaf, especially one associated with a reproductive structure such as a flower, inflorescence axis or cone scale. Bracts are usually different from foliage leaves. They may be smaller, larger, or o ...
subtends each flower; in some species, the bract is large and showy. The
calyx Calyx or calyce (plural "calyces"), from the Latin ''calix'' which itself comes from the Ancient Greek ''κάλυξ'' (''kálux'') meaning "husk" or "pod", may refer to: Biology * Calyx (anatomy), collective name for several cup-like structures ...
usually has four or five lobes; the
corolla Corolla may refer to: *Corolla (botany), the petals of a flower, considered as a unit *Toyota Corolla, an automobile model name *Corolla (headgear) A ''corolla'' is an ancient headdress in the form of a small circlet or crown.stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the fila ...
s number either two or four, arranged in pairs and inserted on the corolla, and the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
is superior and bicarpellated, with axile
placentation Placentation refers to the formation, type and structure, or arrangement of the placenta. The function of placentation is to transfer nutrients, respiratory gases, and water from maternal tissue to a growing embryo, and in some instances to remo ...
. The
fruit In botany, a fruit is the seed-bearing structure in flowering plants that is formed from the ovary after flowering. Fruits are the means by which flowering plants (also known as angiosperms) disseminate their seeds. Edible fruits in partic ...
is a two-celled capsule,
dehiscing Dehiscence is the splitting of a mature plant structure along a built-in line of weakness to release its contents. This is common among fruits, anthers and sporangia. Sometimes this involves the complete detachment of a part; structures that ...
somewhat explosively. In most species, the
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiosper ...
s are attached to a small, hooked stalk (a modified funiculus called a jaculator or a retinaculum) that ejects them from the capsule. This trait is shared by all members of the clade Acanthoideae. A 1995 study of seed expulsion in Acanthaceae used high speed video pictures to show that retinacula propel seeds away from the parent plant when the fruits dehisce, thereby helping the plant gain maximum
seed dispersal In Spermatophyte plants, seed dispersal is the movement, spread or transport of seeds away from the parent plant. Plants have limited mobility and rely upon a variety of dispersal vectors to transport their seeds, including both abiotic vector ...
range. A species well known to
temperate In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout ...
gardeners is bear's breeches (''
Acanthus mollis ''Acanthus mollis'', commonly known as bear's breeches, sea dock, bear's foot plant, sea holly, gator plant or oyster plant, is a species of plant in the family Acanthaceae and is native to the Mediterranean region. It is a leafy, clump-forming ...
''), a herbaceous
perennial plant A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wide ...
with big leaves and flower spikes up to 2 m tall.
Tropical The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
genera familiar to gardeners include '' Thunbergia'' and '' Justicia''. '' Avicennia'', a genus of
mangrove A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in severa ...
trees, usually placed in
Verbenaceae The Verbenaceae ( ), the verbena family or vervain family, is a family of mainly tropical flowering plants. It contains trees, shrubs, and herbs notable for heads, spikes, or clusters of small flowers, many of which have an aromatic smell. The ...
or in its own family, Avicenniaceae, is included in Acanthaceae by the Angiosperm Phylogeny Group on the basis of
molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...
studies that show it to be associated with this family.


Medicinal uses

Traditionally the most important part use in Acanthaceae is the leaves and they are used externally for wounds. Some research has indicated that Acanthaceae possess
antifungal An antifungal medication, also known as an antimycotic medication, is a pharmaceutical fungicide or fungistatic used to treat and prevent mycosis such as athlete's foot, ringworm, candidiasis (thrush), serious systemic infections such as ...
,
cytotoxic Cytotoxicity is the quality of being toxic to cells. Examples of toxic agents are an immune cell or some types of venom, e.g. from the puff adder (''Bitis arietans'') or brown recluse spider (''Loxosceles reclusa''). Cell physiology Treating c ...
,
anti-inflammatory Anti-inflammatory is the property of a substance or treatment that reduces inflammation or swelling. Anti-inflammatory drugs, also called anti-inflammatories, make up about half of analgesics. These drugs remedy pain by reducing inflammation as o ...
, anti-pyretic,
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...
,
insecticidal Insecticides are substances used to kill insects. They include ovicides and larvicides used against insect eggs and larvae, respectively. Insecticides are used in agriculture, medicine, industry and by consumers. Insecticides are claimed t ...
, hepatoprotective, immunomodulatory, anti-platelet aggregation and anti-viral potential. For instance, '' Acanthus ilicifolius'', whose chemical composition has been heavily researched, is widely used in ethnopharmaceutical applications, including in Indian and Chinese
traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...
. Various parts of ''Acanthus ilicifolius'' have been used to treat
asthma Asthma is a long-term inflammatory disease of the airways of the lungs. It is characterized by variable and recurring symptoms, reversible airflow obstruction, and easily triggered bronchospasms. Symptoms include episodes of wheezing, co ...
,
diabetes Diabetes, also known as diabetes mellitus, is a group of metabolic disorders characterized by a high blood sugar level ( hyperglycemia) over a prolonged period of time. Symptoms often include frequent urination, increased thirst and increased ...
,
leprosy Leprosy, also known as Hansen's disease (HD), is a long-term infection by the bacteria '' Mycobacterium leprae'' or '' Mycobacterium lepromatosis''. Infection can lead to damage of the nerves, respiratory tract, skin, and eyes. This nerve d ...
,
hepatitis Hepatitis is inflammation of the liver tissue. Some people or animals with hepatitis have no symptoms, whereas others develop yellow discoloration of the skin and whites of the eyes ( jaundice), poor appetite, vomiting, tiredness, abdominal ...
,
snake bites A snakebite is an injury caused by the bite of a snake, especially a venomous snake. A common sign of a bite from a venomous snake is the presence of two puncture wounds from the animal's fangs. Sometimes venom injection from the bite may occu ...
, and
rheumatoid arthritis Rheumatoid arthritis (RA) is a long-term autoimmune disorder that primarily affects joints. It typically results in warm, swollen, and painful joints. Pain and stiffness often worsen following rest. Most commonly, the wrist and hands are inv ...
. The leaves of '' Acanthus ebracteatus'', noted for their
antioxidant Antioxidants are compounds that inhibit oxidation, a chemical reaction that can produce free radicals. This can lead to polymerization and other chain reactions. They are frequently added to industrial products, such as fuels and lubrica ...
properties, are used for making Thai herbal tea in
Thailand Thailand ( ), historically known as Siam () and officially the Kingdom of Thailand, is a country in Southeast Asia, located at the centre of the Indochinese Peninsula, spanning , with a population of almost 70 million. The country is b ...
and
Indonesia Indonesia, officially the Republic of Indonesia, is a country in Southeast Asia and Oceania between the Indian and Pacific oceans. It consists of over 17,000 islands, including Sumatra, Java, Sulawesi, and parts of Borneo and New Gui ...
.


Phytochemistry

Phytochemical reports on family Acanthaceae are
glycosides In chemistry, a glycoside is a molecule in which a sugar is bound to another functional group via a glycosidic bond. Glycosides play numerous important roles in living organisms. Many plants store chemicals in the form of inactive glycosides. ...
,
flavonoids Flavonoids (or bioflavonoids; from the Latin word ''flavus'', meaning yellow, their color in nature) are a class of polyphenolic secondary metabolites found in plants, and thus commonly consumed in the diets of humans. Chemically, flavonoids ...
, benzonoids,
phenolic compounds In organic chemistry, phenols, sometimes called phenolics, are a class of chemical compounds consisting of one or more hydroxyl groups (— O H) bonded directly to an aromatic hydrocarbon group. The simplest is phenol, . Phenolic compounds are c ...
, naphthoquinone and triterpenoids.


Overview of systematics

Since the first comprehensive classification of Acanthaceae in 1847 by Nees, there have been a few major revisions presented since for the whole family.
Lindau Lindau (german: Lindau (Bodensee), ''Lindau am Bodensee''; ; Low Alemannic German, Low Alemannic: ''Lindou'') is a major Town#Germany, town and Lindau (island), island on the eastern side of Lake Constance (''Bodensee'' in German) in Bavaria, Ge ...
, in 1895, divided the family into the subfamilies Mendoncioideae, Thunbergioideae, Nelsonioideae, and Acanthoideae. Critically, Mendoncioideae, Thunbergioideae, and Nelsonioideae do not possess retinaculate fruits—and it is this distinction, between classifying Acanthaceae into a family that includes those clades with non-retinaculate fruits and one that excludes them, that still persists to the modern day. Bremekamp, in 1965, presented a classification of Acanthaceae that differed from that of Lindau, for his Acanthaceae excluded genera that lack retinaculate fruits. He placed Nelsonioideae within
Scrophulariaceae The Scrophulariaceae are a family of flowering plants, commonly known as the figwort family. The plants are annual and perennial herbs, as well as shrubs. Flowers have bilateral (zygomorphic) or rarely radial (actinomorphic) symmetry. The Scr ...
, classified Thunbergiaceae and Mendonciaceae as distinct families and divided his Acanthaceae into two groups (Acanthoideae and Ruelloideae) based on the presence or absence of
cystolith Cystolith (Gr. "cavity" and "stone") is a botanical term for outgrowths of the epidermal cell wall, usually of calcium carbonate, formed in a cellulose matrix in special cells called lithocysts, generally in the leaf of plants. Cystoliths are pr ...
s, articulate stems, monothecate
anthers The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filam ...
, and colpate
pollen Pollen is a powdery substance produced by seed plants. It consists of pollen grains (highly reduced microgametophytes), which produce male gametes (sperm cells). Pollen grains have a hard coat made of sporopollenin that protects the gametop ...
. In Scotland and Vollesen's 2000 study, they accepted 221
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
and detailed five major groups within Acanthaceae s.s. (that is, those possessing retinaculate fruits), which is equivalent to Acanthoideae Link sensu Lindau 1895. Out of those 221 genera, they placed 201 of them into seven infrafamilial taxa of Acanthaceae, leaving only 20 unplaced. In the current understanding of Acanthaceae, Acanthaceae
s.s. The ''Schutzstaffel'' (SS; also stylized as ''ᛋᛋ'' with Armanen runes; ; "Protection Squadron") was a major paramilitary organization under Adolf Hitler and the Nazi Party in Nazi Germany, and later throughout German-occupied Europe duri ...
includes only those
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English ter ...
s with retinaculate fruits (that is, Acantheae, Barlerieae, Andrographideae, Whitfieldeae, Ruellieae, and Justiceae), while Acanthaceae s.l. includes those clades as well as Thunbergioideae, Nelsonioideae, and '' Avicennia.''


Dating the Acanthaceae lineage

Much research, using both molecular data and
fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserved ...
s, has been conducted in recent years regarding the dating and distribution of the Acanthaceae and Lamiales lineage, although there still remains some ambiguity. In a 2004 study on the molecular phylogenetic dating of asterid flowering plants, researchers estimated 106 million years (MY) for the stem lineage of Lamiales, 67 MY for the stem lineage of Acanthaceae, and 54 MY for the crown node of Acanthaceae (that is, the age of extant lineages with the family). These estimates are older than those based on fossils that can confidently be assigned to Lamiales, which are middle
Eocene The Eocene ( ) Epoch is a geological epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period in the modern Cenozoic Era. The name ''Eocene'' comes from the Ancient Greek (''ēṓs'', ...
in age, ca. 48-37 MY.
Palynomorphs Palynology is the "study of dust" (from grc-gre, παλύνω, palynō, "strew, sprinkle" and ''-logy'') or of "particles that are strewn". A classic palynologist analyses particulate samples collected from the air, from water, or from deposit ...
that definitively show the existence of Acanthaceae are known from the upper
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
, with the oldest ca. 22 MY.


Selected genera

, the Germplasm Resources Information Network accepts 217 genera, while 206 accepted genera are listed by
Plants of the World Online Plants of the World Online (POWO) is an online database published by the Royal Botanic Gardens, Kew. It was launched in March 2017 with the ultimate aim being "to enable users to access information on all the world's known seed-bearing plants by ...
(the Royal Botanic Gardens, Kew).


Excluded genera

* '' Thomandersia'' Baill.Thomandersiaceae


References

*


External links


Tree of Life Acanthaceae


Flowers in Israel.

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{{Taxonbar, from=Q53475 Lamiales families