Dwarfing is a process in which a breed of animals or
cultivar
A cultivar is a type of cultivated plant that people have selected for desired traits and when propagated retain those traits. Methods used to propagate cultivars include: division, root and stem cuttings, offsets, grafting, tissue culture ...
of plants is changed to become significantly smaller than standard members of their species. The effect can be induced through human intervention or non-human processes, and can include genetic, nutritional or hormonal means. Used most specifically, dwarfing includes pathogenic changes in the structure of an organism (for example, the bulldog, a genetically
achondroplastic
Achondroplasia is a genetic disorder with an autosomal dominant pattern of inheritance whose primary feature is dwarfism. In those with the condition, the arms and legs are short, while the torso is typically of normal length. Those affected ...
dog breed), in contrast to non-pathogenic proportional reduction in stature (such as the
whippet
The Whippet is a dog breed of medium size. It is a sighthound breed that originated in England, descended from the Greyhound. Whippets today still strongly resemble a smaller Greyhound. Part of the hound group, Whippets have relatively ...
, a small sighthound dog breed).
Animals
In animals, including humans, dwarfism has been described in several ways. Shortened stature can result from
growth hormone deficiency
Growth hormone deficiency (GHD), or human growth hormone deficiency, is a medical condition resulting from not enough growth hormone (GH). Generally the most noticeable symptom is that an individual attains a short height. Newborns may also prese ...
,
starvation
Starvation is a severe deficiency in caloric energy intake, below the level needed to maintain an organism's life. It is the most extreme form of malnutrition. In humans, prolonged starvation can cause permanent organ damage and eventually, dea ...
,
portal systemic shunts
Portal often refers to:
*Portal (architecture), an opening in a wall of a building, gate or fortification, or the extremities (ends) of a tunnel
Portal may also refer to:
Arts and entertainment Gaming
* ''Portal'' (series), two video games ...
,
renal disease
Kidney disease, or renal disease, technically referred to as nephropathy, is damage to or disease of a kidney. Nephritis is an inflammatory kidney disease and has several types according to the location of the inflammation. Inflammation can b ...
,
hypothyroidism
Hypothyroidism (also called ''underactive thyroid'', ''low thyroid'' or ''hypothyreosis'') is a disorder of the endocrine system in which the thyroid gland does not produce enough thyroid hormone. It can cause a number of symptoms, such as ...
diabetes mellitus
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 ...
and other conditions. Any of these conditions can be established in a population through
genetic engineering
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including ...
,
selective breeding
Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
, or
insular dwarfism
Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is dist ...
, or some combination of the above.
Dwarfing can produce more practical breeds that can fit in small accommodations, or may appeal aesthetically, as well as other associated side effects. Smaller stature may be a deliberate goal of breeding programs, or it may be a side effect of other breeding goals.
Nonpurposeful dwarfing
In some husbandry conditions, humans created dwarf breeds, or allowed them to develop, without specifically selecting for smaller animals. It is likely that the
Shetland sheep
The Shetland is a small, wool-producing breed of sheep originating in the Shetland Isles, Scotland, but is now also kept in many other parts of the world. It is part of the Northern European short-tailed sheep group, and it is closely related ...
breed, Shetland collie breed of dogs, and various pony breeds of horses developed in this manner. In the case of the Shetland sheep and collies, it is likely that environmental conditions, such as a lack of abundant fodder, led to farmers selecting smaller animals who continued to reproduce on limited food over larger animals who did not reproduce well on limited diets. In this case, the emphasis was on selecting for survival and reproduction, not size.
Purposeful dwarfing
Humans have encouraged the deliberate development of dwarf breeds of many domestic animals, including horses, cattle, dogs, and chickens. Some have been breeds of smaller animals that were not originally selected for size, but are now held to specific sizes by a breed standard. In many cases, the exact physiological mechanism that alters the growth of individuals in that breed is not well known, and some breeds have multiple mechanisms at play.
As the genetic mutations that cause dwarfing occur in many species, dwarf animals can be the offspring of normal-appearing animals. Even in breeds which have not been selected for dwarfing, some genetic lines may show a tendency to produce dwarfs, which may be encouraged by deliberate breeding. This often takes the form of in-breeding to concentrate recessive genes, and can result in other genetic abnormalities being established in the population.
Some animal breeds that have been formally subject to dwarfing include:
* Rabbits ( American Fuzzy Lop, Britannia Petite (US), Dwarf Hotot, Florida White Rabbit, Jersey Wooly, Lionhead
Polish
Polish may refer to:
* Anything from or related to Poland, a country in Europe
* Polish language
* Poles, people from Poland or of Polish descent
* Polish chicken
*Polish brothers (Mark Polish and Michael Polish, born 1970), American twin screenwr ...
(UK),
Holland Lop
Holland Lop is a breed of rabbit that was recognized by the American Rabbit Breeders Association (ARBA) in 1979 and by the Netherlands' Governing Rabbit Council in 1984. The Holland Lop, with a maximum weight of (as stipulated by ARBA), is one ...
Netherland Dwarf
The Netherland Dwarf is a breed of domestic rabbit that originated in the Netherlands. Weighing , the Netherland Dwarf is one of the smallest rabbit breeds. Its popularity as a pet or show rabbit may stem from its neotenic appearance. The Nethe ...
English Bulldog
The Bulldog is a British breed of dog of mastiff type. It may also be known as the English Bulldog or British Bulldog. It is of medium size, a muscular, hefty dog with a wrinkled face and a distinctive pushed-in nose.French Bulldog, Chihuahua, Corgi,
Whippet
The Whippet is a dog breed of medium size. It is a sighthound breed that originated in England, descended from the Greyhound. Whippets today still strongly resemble a smaller Greyhound. Part of the hound group, Whippets have relatively ...
Nigerian Dwarf
The Nigerian Dwarf is an American breed of dwarf goat. Like the American Pygmy Goat, it derives from the West African Dwarf group of breeds of West Africa.
History
Between about 1930 and 1960 a variety of small goats of the West African ...
,
Pygmy Goat
The American Pygmy is an American breed of achondroplastic goat. It is small, compact and stockily built. Like the Nigerian Dwarf, it derives from the West African Dwarf group of breeds of West Africa. Between 1930 and 1960, animals of this t ...
)
* Cattle (Dwarf Brahman,
Lowline cattle
The Australian Lowline is a modern Australian breed of small, polled beef cattle. It was the result of a selective breeding experiment using black Aberdeen Angus cattle at the Agricultural Research Centre of the Department of Agriculture of New ...
Guinea Cracker Cattle,
Dexter cattle
The Dexter is an Irish breed of small cattle. It originated in the eighteenth century in County Kerry, in south-western Ireland, and appears to be named after a man named Dexter, who was factor of the estates of Lord Hawarden on Valentia Isla ...
)
* Equines (Miniature Donkey,
Fell Pony
The Fell pony is a versatile working breed of mountain and moorland pony originating on Cumberland and Westmorland farms of northwest England and is used for riding and driving. The breed is closely related to its geographic neighbour, the ...
,
Hackney Pony
The Hackney pony is a breed of pony closely related to the Hackney horse. Originally bred to pull carriages, they are used today primarily as show ponies. The breed does not have its own stud book, but shares one with the Hackney horse in ...
Miniature horse
A miniature horse is a breed or type of horse characterised by its small size. Usually it has been bred to display in miniature the physical characteristics of a full-sized horse, but to be little over in height, or even less. Although such hor ...
,
Shetland Pony
The Shetland pony is a Scottish breed of pony originating in the Shetland Isles in the north of Scotland. It may stand up to at the withers. It has a heavy coat and short legs, is strong for its size, and is used for riding, driving, and p ...
, others)
Plants
As with animals, plants can be dwarfed through
genetic engineering
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including ...
and
selective breeding
Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
, but can also undergo natural, morphological changes to acclimatize to environmental stresses such as soil quality,Hutchings, M. J.; de Kroon, H. (1994), "Foraging in Plants: The Role of Morphological Plasticity in Resource Acquisition", ''Adv. Ecol. Res.''
25: 159–238 light, drought, flood,Else, M. A.; et al. (1996), "Stomatal Closure in Flooded Tomato Plants Involves Abscisic Acid and a Chemically Unidentified Anti-Transpirant in Xylem Sap", ''Plant Physiol'' 112: 239-247 cold,Okamoto, T.; Tsurumi, S.; Shibasaki, K.; Obana, Y.; Takaji, H.; Oono Y.; Rahman, A. (2008), "Genetic Dissection of Hormonal Responses in the Roots of ''Arabidopsis'' Grown Under Continuous Mechanical Impedance", ''Plant
Physiol.'' 146: 1651–1662 infection,Scholthof, H. B.; Scholthof, K. B. G.; Jackson, A. O. (1995), "Identification of Tomato Bushy Stunt Virus Host-Specific Symptom Determinants by Expression of Individual Genes from a Potato Virus X Vector", ''Plant Cell'' 7: 1157-1172 and
herbivory
A herbivore is an animal anatomically and physiologically adapted to eating plant material, for example foliage or marine algae, for the main component of its diet. As a result of their plant diet, herbivorous animals typically have mouthp ...
Chouinard, A.; Filion, L. (2005), "Impact of Introduced White-Tailed Deer and Native Insect Defoliators on the Density and Growth of Conifer Saplings on Anticosti Island, Quebec", ''Ecoscience'' 12: 506-518 resulting in a dwarfed stature. Plants dwarfed due to environmental stress are said to be "
stunted
Stunted growth is a reduced growth rate in human development. It is a primary manifestation of malnutrition (or more precisely undernutrition) and recurrent infections, such as diarrhea and helminthiasis, in early childhood and even before birth, ...
." The majority of dwarfing in plants occurs not from the damage environmental stresses inflict on them, but instead by hormones produced in response to the stress.Zhang, Y.; Turner, J. G. (2008), "Wound-Induced Endogenous Jasmonates Stunt Plant Growth by Inhibiting Mitosis", ''PLoS ONE'' 3: e3699
Plant hormone
Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
s act as a signal to the various tissues of plants inducing one or more responses, the class of
plant hormone
Plant hormone (or phytohormones) are signal molecules, produced within plants, that occur in extremely low concentrations. Plant hormones control all aspects of plant growth and development, from embryogenesis, the regulation of organ size, pa ...
responsible for dwarfing in plants due to injury are called
jasmonate
Jasmonate (JA) and its derivatives are lipid-based plant hormones that regulate a wide range of processes in plants, ranging from growth and photosynthesis to reproductive development. In particular, JAs are critical for plant defense against her ...
s. Such responses include, but are not limited to: less frequent cell divisions and reduction of cell elongation.Swarup, R.; Perry, P.; Hagenbeek, D.; Van Der Straeten, D.; Beemster, G. T. S.; et al. (2007), "Ethylene Upregulates Auxin Biosynthesis in ''Arabidopsis'' Seedlings to Enhance Inhibition of Root Cell Elongation", ''Plant Cell'' 19: 2186–2196
Dwarfing trees
In
horticulture
Horticulture is the branch of agriculture that deals with the art, science, technology, and business of plant cultivation. It includes the cultivation of fruits, vegetables, nuts, seeds, herbs, sprouts, mushrooms, algae, flowers, seaweeds and no ...
, dwarfing can be considered a desirable characteristic in modern
orchard
An orchard is an intentional plantation of trees or shrubs that is maintained for food production. Orchards comprise fruit- or nut-producing trees which are generally grown for commercial production. Orchards are also sometimes a feature of la ...
s. This kind of dwarfing can be attained through
selective breeding
Selective breeding (also called artificial selection) is the process by which humans use animal breeding and plant breeding to selectively develop particular phenotypic traits (characteristics) by choosing which typically animal or plant ma ...
,
genetic engineering
Genetic engineering, also called genetic modification or genetic manipulation, is the modification and manipulation of an organism's genes using technology. It is a set of technologies used to change the genetic makeup of cells, including ...
, or more often, scions are grafted on to dwarfing rootstocks. Almost all modern
apple
An apple is an edible fruit produced by an apple tree (''Malus domestica''). Apple trees are cultivated worldwide and are the most widely grown species in the genus '' Malus''. The tree originated in Central Asia, where its wild ances ...
s in commercial use are propagated as dwarf or semi-dwarf trees for ease of picking and spraying.
Dwarfing fruit trees acts through a reduction in the nutrients which travel from the roots through the trunk to the leaves and buds. Many commercial orchards of various species use this technique to improve the overall health and productivity of the individual trees. An individual tree may be made up of three or more separate cultivars - one for the root system, which is generally selected for good stability and resistance to soil-borne diseases, one for the trunk, which modifies the overall height of the tree, and one for the productive limbs and buds, which actually produces the fruit. Frequently, the root system stock is the most resistant to cold damage - both by natural selection and by protection from the cold air by the earth. When frost severely damages a tree, the more productive branch and bud cultivar may be killed off, leaving the root to sprout new stalks. In the case of oranges and other citrus, this results in sweet orange trees being frozen back so that the more hardy, cold-tolerant sour orange rootstock puts out new growth.
Dwarfing grains
Dwarfing genes are widely used in creating more productive food plants, such as grains. One condition that results in loss of grain crops is called 'lodging', where heavy ears of almost ripe grain bend the stalk until the grain touches the ground, becomes wet, and spoils. During the
Green Revolution
The Green Revolution, also known as the Third Agricultural Revolution, was a period of technology transfer initiatives that saw greatly increased crop yields and agricultural production. These changes in agriculture began in developed countrie ...
, research that identified wheat reduced-height genes (''Rht'') and a rice semidwarf gene (''sd1'') resulted in crops that yielded significantly more harvestable grain.
Dwarfism in chickens
Dwarfism in chickens is an inherited condition found in chickens consisting of a significant delayed growth, resulting in adult individuals with a distinctive small size in comparison with normal specimens of the same breed or population.
The af ...
*
Fruit tree propagation
Fruit tree propagation is usually carried out vegetatively (non-sexually) by grafting or budding a desired variety onto a suitable rootstock.
Perennial plants can be propagated either by sexual or vegetative means. Sexual reproduction begins wh ...
*
Genomics of domestication
Domesticated species and the human populations that domesticate them are typified by a mutualistic relationship of interdependence, in which humans have over thousands of years modified the genomics of domesticated species. Genomics is the study o ...
*
Green revolution
The Green Revolution, also known as the Third Agricultural Revolution, was a period of technology transfer initiatives that saw greatly increased crop yields and agricultural production. These changes in agriculture began in developed countrie ...
Deep-sea gigantism
In zoology, deep-sea gigantism or abyssal gigantism is the tendency for species of invertebrates and other deep-sea dwelling animals to be larger than their shallower-water relatives across a large taxonomic range. Proposed explanations for thi ...
Insular dwarfism
Insular dwarfism, a form of phyletic dwarfism, is the process and condition of large animals evolving or having a reduced body size when their population's range is limited to a small environment, primarily islands. This natural process is dist ...
*
Island gigantism
Island gigantism, or insular gigantism, is a biological phenomenon in which the size of an animal species isolated on an island increases dramatically in comparison to its mainland relatives. Island gigantism is one aspect of the more general Fos ...
*
Prostrate shrub
A prostrate shrub is a woody plant, most of the branches of which lie upon or just above the ground, rather than being held erect as are the branches of most trees and shrubs.
Background
Prostration may occur because the supporting tissues in ste ...