Phyllody

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Phyllody on a purple coneflower (Echinacea purpurea) Phyllody on Coneflower with aster yellows.jpg
Phyllody on a purple coneflower ( Echinacea purpurea )

Phyllody is the abnormal development of floral parts into leafy structures. It is generally caused by phytoplasma or virus infections, [1] though it may also be because of environmental factors that result in an imbalance in plant hormones. [2] Phyllody causes the affected plant to become partially or entirely sterile, as it is unable to produce normal flowers. [1] [3]

Contents

The condition is also known as phyllomorphy or frondescence; [4] though the latter may sometimes refer more generically to foliage, leafiness, or the process of leaf growth. Phyllody is usually differentiated from floral virescence, wherein the flowers merely turn green in color but otherwise retain their normal structure. [5] However, floral virescence and phyllody (along with witch's broom and other growth abnormalities), commonly occur together as symptoms of the same diseases. [6] The term chloranthy is also often used for phyllody (particularly flowers exhibiting complete phyllody, such that it resembles leaf buds more than flowers), though in some cases it may refer to floral virescence. [7]

History

In the late 18th century, the German poet and philosopher Johann Wolfgang von Goethe noted strange-looking rose flowers where the flower organs were replaced by leafy or stem-like structures. This led him to hypothesize that plant organs arising from the stem are simply modifications of the same basic leaf organ. During growth, these organs naturally differentiate into specialized or generalized structures like petals or leaves. However, if certain factors interfere during the early growth stages, these organs can develop into something other than the original "plan of construction". He called this abnormal growth "metamorphosis" and it is the main topic of his essay Versuch die Metamorphose der Pflanzen zu erklären (1790), better known in English as the Metamorphosis of Plants . [2] [5] Goethe's hypothesis was poorly received by other scientists during his time, but it is now known to be essentially correct. The concepts he discusses while describing metamorphosis is now known as homology, the basis of the modern science of comparative anatomy and a discovery that is usually credited to the English biologist Sir Richard Owen. [5] [8]

In 1832, the German-American botanist George Engelmann described the same condition in his work De Antholysi Prodromus. He gave it the name "frondescence". [5] Nineteen years later, the Belgian botanist Charles Jacques Édouard Morren also investigated the phenomenon in his book Lobelia (1851). Morren called the condition "phyllomorphy", and unlike Engelmann, Morren explicitly distinguished phyllomorphy (wherein the floral parts are replaced by leaf-like structures) from virescence (wherein the affected parts, not necessarily floral, turn green but retain the original form or structure). [5]

The term "phyllody" was coined by the English botanist Maxwell T. Masters in his book on plant abnormalities, Vegetable Teratology (1869). [2] The term is derived from Scientific Latin phyllodium, which is itself derived from Ancient Greek φυλλώδης (phullodes, 'leaf-like'). [9] [10] Like Morren, Masters also distinguished phyllody from virescence. He acknowledged "frondescence" and "phyllomorphy" as synonyms of phyllody. [5]

Description

Phyllody is characterized by the partial or complete replacement of floral organs with true leaves. Phyllody can affect bracts, the calyx (sepals), corolla (petals), the gynoecium (carpels/pistils), and the androecium (stamens). Phyllody may be partial, affecting only some sets of floral organs or even only half of a set of floral organs (e.g. only three petals out of six in a single flower); or it can be complete, with all the floral organs replaced by leaves. [5]

Illustration from Vegetable Teratology (1869), showing a Petunia flower with stamens partially replaced by "stalked" leaves. The stalks are actually the retained filaments of the stamens, while the anthers have been replaced by small leaves. Phyllody - Flower of Petunia, opened to show the stamens partially replaced by stalked leaves.jpg
Illustration from Vegetable Teratology (1869), showing a Petunia flower with stamens partially replaced by "stalked" leaves. The stalks are actually the retained filaments of the stamens, while the anthers have been replaced by small leaves.
Phyllody 1.jpg
Longitudinal section of a rose flower exhibiting phyllody. Despite the apparent hips, the reproductive organs are completely absent and have been replaced by leaves.
Rose hip 02 ies.jpg
Longitudinal section of a normal developing rose hip

Phyllody of the bracts is common among plants which bear catkin (amentaceous) inflorescences. They are very common among members of the genus Plantago , for example, as well as the common hop ( Humulus lupulus ). Involucral bracts of the flowers of members of the family Asteraceae like dahlias and dandelions, may also be affected. [5]

Sepals that exhibit phyllody are usually hard to detect due to fact that most sepals already resemble leaves. Close examination, however, can reveal differences in venation in normal sepals and sepals that exhibit phyllody. The full development of perfect leaves from sepals is more common among flowers that have united sepals (monosepalous) than in flowers with separated sepals (polysepalous). [5]

Phyllody of the petals can be expressed more mildly as a simple change in shape and color (in which case, it's more accurately virescence), or it can be expressed as fully formed leaves. It is more common among flowers which exhibit corollas of distinct petals (polypetalous) than in flowers in which the petals are fused into a single tube or bowl-like structure (monopetalous). [5]

Phyllody of the stamens is rare. In fact, the stamens are the least likely of the floral organs to be affected by phyllody. This is thought to be because the stamens are the most highly differentiated organs in flowers. [5]

In contrast, phyllody of the carpels is much more common than the corresponding changes in stamens. Usually, phyllody affects the proximal parts of the carpel (the ovary) more than the distal parts (the style and stigma). The ovule itself may be exposed on the edges or on the inner surface of the carpel if the ovary becomes leaf-like. If the ovule is affected by phyllody, it develops separately from the rest of the carpel. The best known example of phyllody of the carpels is found in the Japanese cherry ( Prunus serrulata ), in which one or both of the carpels can become leaf-like (although the distal half of the style and the stigma are usually unaffected). Incidentally, some Japanese cherry cultivars also exhibit "doubling" of the petals due to petalody, where a second corolla develops instead of stamens. [5] [11]

Phyllody on a Brinjal (Solanum melongena) Brinjal phyllody.jpg
Phyllody on a Brinjal ( Solanum melongena )

Causes

Biotic

In many cultivated plants, phyllody is caused by infections of plant pathogens and/or infestations of ectoparasites. Aside from exhibiting phyllody, they may also exhibit other symptoms like virescence, witch's brooms, chlorosis, and stunted growth. [2] [6] [12] Examples of these biotic factors include:

In addition to causing phyllody itself, insects and other ectoparasites also serve as disease vectors that can spread phyllody to other nearby plants. [19] The most common of these insect vectors are leafhoppers, [19] an example of which is Hishimonus phycitis , which transmits the phytoplasma-caused little leaf phyllody in eggplants. [20] The broken-backed bug ( Taylorilygus apicalis ) is another insect vector of a phytoplasma-caused phyllody in species of Parthenium . [21] Other ectoparasite vectors include eriophyid mites, like the rose leaf curl mite ( Phyllocoptes fructiplilus ) which is known to be the primary vector of the rose rosette disease; [19] [22] and the chrysanthemum rust mite ( Paraphytoptus chrysanthemi ) which transmits phytoplasma-caused phyllody in species of chrysanthemums. [23] [24]

Abiotic

Environmental abiotic factors like hot weather or water stress that result in an imbalance in plant hormones during flowering can cause phyllody. These can usually be differentiated from phyllody caused by biotic factors by the simultaneous presence of healthy and abnormal flowers. When conditions normalize, the plants resume normal flowering. The susceptibility of plants to environmentally caused phyllody can be genetic. [2]

Artificial

Phyllody can be artificially induced by applying cytokinins (CK), plant hormones responsible for cell division, as well as apical dominance and axillary bud growth. Conversely, it can be subsequently suppressed with the application of gibberellins (GA), plant hormones responsible for stem elongation, flowering, and sex expression. [2]

Other related floral development abnormalities are:

Phyllody in plant breeding

Phyllody in the green rose (Rosa chinensis 'Viridiflora') Green rose 3.jpg
Phyllody in the green rose ( Rosa chinensis 'Viridiflora')

In some cases, the occurrence of phyllody has been utilized in plant breeding. One of the most well known examples is the green rose ( Rosa chinensis 'Viridiflora'), an ancient Chinese rose cultivar which exhibits green leafy bracts in tight flower-like clusters. In green rose, artificial selection has enabled phyllody to be expressed as a stable mutation. [2] [25] [26]

See also

Related Research Articles

<span class="mw-page-title-main">Petal</span> Part of most types of flower

Petals are modified leaves that surround the reproductive parts of flowers. They are often brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the corolla. Petals are usually accompanied by another set of modified leaves called sepals, that collectively form the calyx and lie just beneath the corolla. The calyx and the corolla together make up the perianth, the non-reproductive portion of a flower. When the petals and sepals of a flower are difficult to distinguish, they are collectively called tepals. Examples of plants in which the term tepal is appropriate include genera such as Aloe and Tulipa. Conversely, genera such as Rosa and Phaseolus have well-distinguished sepals and petals. When the undifferentiated tepals resemble petals, they are referred to as "petaloid", as in petaloid monocots, orders of monocots with brightly colored tepals. Since they include Liliales, an alternative name is lilioid monocots.

<span class="mw-page-title-main">Stamen</span> Male organ of a flower

The stamen is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium.

<i>Trillium grandiflorum</i> Species of flowering plant

Trillium grandiflorum, the white trillium, large-flowered trillium, great white trillium, white wake-robin or French: trille blanc, is a species of flowering plant in the family Melanthiaceae. A monocotyledonous, herbaceous perennial, the plant is native to eastern North America, from northern Quebec to the southern parts of the United States through the Appalachian Mountains into northernmost Georgia and west to Minnesota. There are also several isolated populations in Nova Scotia, Maine, southern Illinois, and Iowa.

<span class="mw-page-title-main">Witch's broom</span> Type of deformity in a woody plant

Witch's broom or witches' broom is a deformity in a woody plant, typically a tree, where the natural structure of the plant is changed. A dense mass of shoots grows from a single point, with the resulting structure resembling a broom or a bird's nest. It is sometimes caused by pathogens.

<i>Phytoplasma</i> Genus of bacteria

Phytoplasmas are obligate intracellular parasites of plant phloem tissue and of the insect vectors that are involved in their plant-to-plant transmission. Phytoplasmas were discovered in 1967 by Japanese scientists who termed them mycoplasma-like organisms. Since their discovery, phytoplasmas have resisted all attempts at in vitro culture in any cell-free medium; routine cultivation in an artificial medium thus remains a major challenge. Phytoplasmas are characterized by the lack of a cell wall, a pleiomorphic or filamentous shape, a diameter normally less than 1 μm, and a very small genome.

<span class="mw-page-title-main">Gynoecium</span> Female organs of a flower

Gynoecium is most commonly used as a collective term for the parts of a flower that produce ovules and ultimately develop into the fruit and seeds. The gynoecium is the innermost whorl of a flower; it consists of pistils and is typically surrounded by the pollen-producing reproductive organs, the stamens, collectively called the androecium. The gynoecium is often referred to as the "female" portion of the flower, although rather than directly producing female gametes, the gynoecium produces megaspores, each of which develops into a female gametophyte which then produces egg cells.

<span class="mw-page-title-main">ABC model of flower development</span> Model for genetics of flower development

The ABC model of flower development is a scientific model of the process by which flowering plants produce a pattern of gene expression in meristems that leads to the appearance of an organ oriented towards sexual reproduction, a flower. There are three physiological developments that must occur in order for this to take place: firstly, the plant must pass from sexual immaturity into a sexually mature state ; secondly, the transformation of the apical meristem's function from a vegetative meristem into a floral meristem or inflorescence; and finally the growth of the flower's individual organs. The latter phase has been modelled using the ABC model, which aims to describe the biological basis of the process from the perspective of molecular and developmental genetics.

<span class="mw-page-title-main">Flower</span> Reproductive structure in flowering plants

A flower, sometimes known as a bloom or blossom, is the reproductive structure found in flowering plants. Flowers produce gametophytes, which in flowering plants consist of a few haploid cells which produce gametes. The "male" gametophyte, which produces non-motile sperm, is enclosed within pollen grains; the "female" gametophyte is contained within the ovule. When pollen from the anther of a flower is deposited on the stigma, this is called pollination. Some flowers may self-pollinate, producing seed using pollen from the same flower or a different flower of the same plant, but others have mechanisms to prevent self-pollination and rely on cross-pollination, when pollen is transferred from the anther of one flower to the stigma of another flower on a different individual of the same species.

<span class="mw-page-title-main">Aster yellows</span> Plant disease

Aster yellows is a chronic, systemic plant disease caused by several bacteria called phytoplasma. The aster yellows phytoplasma (AYP) affects 300 species in 38 families of broad-leaf herbaceous plants, primarily in the aster family, as well as important cereal crops such as wheat and barley. Symptoms are variable and can include phyllody, virescence, chlorosis, stunting, and sterility of flowers. The aster leafhopper vector, Macrosteles quadrilineatus, moves the aster yellows phytoplasma from plant to plant. Its economic burden is primarily felt in the carrot crop industry, as well as the nursery industry. No cure is known for plants infected with aster yellows. Infected plants should be removed immediately to limit the continued spread of the phytoplasma to other susceptible plants. However, in agricultural settings such as carrot fields, some application of chemical insecticides has proven to minimize the rate of infection by killing the vector.

Merosity refers to the number of component parts in a distinct whorl of a plant structure. The term is most commonly used in the context of a flower where it refers to the number of sepals in a whorl of the calyx, the number of petals in a whorl of the corolla, the number of stamens in a whorl of the androecium, or the number of carpels in a whorl of the gynoecium. The term may also be used to refer to the number of leaves in a leaf whorl.

This page provides a glossary of plant morphology. Botanists and other biologists who study plant morphology use a number of different terms to classify and identify plant organs and parts that can be observed using no more than a handheld magnifying lens. This page provides help in understanding the numerous other pages describing plants by their various taxa. The accompanying page—Plant morphology—provides an overview of the science of the external form of plants. There is also an alphabetical list: Glossary of botanical terms. In contrast, this page deals with botanical terms in a systematic manner, with some illustrations, and organized by plant anatomy and function in plant physiology.

Important structures in plant development are buds, shoots, roots, leaves, and flowers; plants produce these tissues and structures throughout their life from meristems located at the tips of organs, or between mature tissues. Thus, a living plant always has embryonic tissues. By contrast, an animal embryo will very early produce all of the body parts that it will ever have in its life. When the animal is born, it has all its body parts and from that point will only grow larger and more mature. However, both plants and animals pass through a phylotypic stage that evolved independently and that causes a developmental constraint limiting morphological diversification.

<span class="mw-page-title-main">Double-flowered</span> Varieties of flowers with extra petals

"Double-flowered" describes varieties of flowers with extra petals, often containing flowers within flowers. The double-flowered trait is often noted alongside the scientific name with the abbreviation fl. pl.. The first abnormality to be documented in flowers, double flowers are popular varieties of many commercial flower types, including roses, camellias and carnations. In some double-flowered varieties all of the reproductive organs are converted to petals — as a result, they are sexually sterile and must be propagated through cuttings. Many double-flowered plants have little wildlife value as access to the nectaries is typically blocked by the mutation.

This glossary of botanical terms is a list of definitions of terms and concepts relevant to botany and plants in general. Terms of plant morphology are included here as well as at the more specific Glossary of plant morphology and Glossary of leaf morphology. For other related terms, see Glossary of phytopathology, Glossary of lichen terms, and List of Latin and Greek words commonly used in systematic names.

<i>Enemion biternatum</i> Species of flowering plant

Enemion biternatum, commonly known as the false rue-anemone, is a spring ephemeral native to moist deciduous woodland in the eastern United States and extreme southern Ontario.

<span class="mw-page-title-main">Whorl (botany)</span> Arrangement of plant parts around the stem

In botany, a whorl or verticil is a whorled arrangement of leaves, sepals, petals, stamens, or carpels that radiate from a single point and surround or wrap around the stem or stalk. A leaf whorl consists of at least three elements; a pair of opposite leaves is not called a whorl.

<span class="mw-page-title-main">Floral formula</span> Notation representing flowers structure

A floral formula is a notation for representing the structure of particular types of flowers. Such notations use numbers, letters and various symbols to convey significant information in a compact form. They may represent the floral form of a particular species, or may be generalized to characterize higher taxa, usually giving ranges of numbers of organs. Floral formulae are one of the two ways of describing flower structure developed during the 19th century, the other being floral diagrams. The format of floral formulae differs according to the tastes of particular authors and periods, yet they tend to convey the same information.

Virescence is the abnormal development of green pigmentation in plant parts that are not normally green, like shoots or flowers. Virescence is closely associated with phyllody and witch's broom. They are often symptoms of the same disease affecting the plants, typically those caused by phytoplasmas. The term chloranthy is also sometimes used for floral virescence, though it is more commonly used for phyllody.

<span class="mw-page-title-main">Floral morphology</span>

In botany, floral morphology is the study of the diversity of forms and structures presented by the flower, which, by definition, is a branch of limited growth that bears the modified leaves responsible for reproduction and protection of the gametes, called floral pieces.

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