Inflorescence

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Aloe hereroensis, showing inflorescence with branched peduncle Aloe hereroensis Auob C15.JPG
Aloe hereroensis , showing inflorescence with branched peduncle
Amorphophallus titanum has the world's largest unbranched inflorescence. Photo of the plant in bloom in 2000 at Fairchild Tropical Botanic Garden in Miami, Florida, US Amorphophallus at Fairchild.jpg
Amorphophallus titanum has the world's largest unbranched inflorescence. Photo of the plant in bloom in 2000 at Fairchild Tropical Botanic Garden in Miami, Florida, US

An inflorescence, in a flowering plant, is a group or cluster of flowers arranged on a stem that is composed of a main branch or a system of branches. [1] An inflorescence is categorized on the basis of the arrangement of flowers on a main axis (peduncle) and by the timing of its flowering (determinate and indeterminate). [2]

Contents

Morphologically, an inflorescence is the modified part of the shoot of seed plants where flowers are formed on the axis of a plant. The modifications can involve the length and the nature of the internodes and the phyllotaxis, as well as variations in the proportions, compressions, swellings, adnations, connations and reduction of main and secondary axes.[ citation needed ]

One can also define an inflorescence as the reproductive portion of a plant that bears a cluster of flowers in a specific pattern.[ citation needed ]

General characteristics

Inflorescences are described by many different characteristics including how the flowers are arranged on the peduncle, the blooming order of the flowers, and how different clusters of flowers are grouped within it. These terms are general representations as plants in nature can have a combination of types. Because flowers facilitate plant reproduction, inflorescence characteristics are largely a result of natural selection. [3]

The stem holding the whole inflorescence is called a peduncle. The main axis (also referred to as major stem) above the peduncle bearing the flowers or secondary branches is called the rachis. The stalk of each flower in the inflorescence is called a pedicel. A flower that is not part of an inflorescence is called a solitary flower and its stalk is also referred to as a peduncle. Any flower in an inflorescence may be referred to as a floret, especially when the individual flowers are particularly small and borne in a tight cluster, such as in a pseudanthium. The fruiting stage of an inflorescence is known as an infructescence. Inflorescences may be simple (single) or complex (panicle). The rachis may be one of several types, including single, composite, umbel, spike or raceme. [ citation needed ]

In some species the flowers develop directly from the main stem or woody trunk, rather than from the plant's main shoot. This is called cauliflory and is found across a number of plant families. [4] An extreme version of this is flagelliflory where long, whip-like branches grow from the main trunk to the ground and even below it. Inflorescences form directly on these branches. [5]

Bracts

Inflorescences usually have modified foliage different from the vegetative part of the plant. Considering the broadest meaning of the term, any leaf associated with an inflorescence is called a bract . A bract is usually located at the node where the main stem of the inflorescence forms, joined to the rachis of the plant, but other bracts can exist within the inflorescence itself. They serve a variety of functions which include attracting pollinators and protecting young flowers. According to the presence or absence of bracts and their characteristics we may distinguish the following:

If many bracts are present and they are strictly connected to the stem, like in the family Asteraceae, the bracts might collectively be called an involucre. If the inflorescence has a second unit of bracts further up the stem, they might be called an involucel.

Terminal flower

Plant organs can grow according to two different schemes, namely monopodial or racemose and sympodial or cymose. In inflorescences these two different growth patterns are called indeterminate and determinate respectively, and indicate whether a terminal flower is formed and where flowering starts within the inflorescence.

Indeterminate and determinate inflorescences are sometimes referred to as open and closed inflorescences respectively. The indeterminate patterning of flowers is derived from determinate flowers. It is suggested that indeterminate flowers have a common mechanism that prevents terminal flower growth. Based on phylogenetic analyses, this mechanism arose independently multiple times in different species. [6]

In an indeterminate inflorescence there is no true terminal flower and the stem usually has a rudimentary end. In many cases the last true flower formed by the terminal bud (subterminal flower) straightens up, appearing to be a terminal flower. Often a vestige of the terminal bud may be noticed higher on the stem.

In determinate inflorescences the terminal flower is usually the first to mature (precursive development), while the others tend to mature starting from the base of the stem. This pattern is called acropetal maturation. When flowers start to mature from the top of the stem, maturation is basipetal, whereas when the central mature first, maturation is divergent.

Phyllotaxis

As with leaves, flowers can be arranged on the stem according to many different patterns. See 'Phyllotaxis' for in-depth descriptions.

Similarly arrangement of leaf in bud is called Ptyxis.

When a single or a cluster of flower(s) is located at the axil of a bract, the location of the bract in relation to the stem holding the flower(s) is indicated by the use of different terms and may be a useful diagnostic indicator.

Typical placement of bracts include:

Metatopic placement of bracts include:

Organization

There is no general consensus in defining the different inflorescences. The following is based on Focko Weberling's Morphologie der Blüten und der Blütenstände (Stuttgart, 1981). The main groups of inflorescences are distinguished by branching. Within these groups, the most important characteristics are the intersection of the axes and different variations of the model. They may contain many flowers (pluriflor) or a few (pauciflor). Inflorescences can be simple or compound.

Simple inflorescences

Inflorescence of sessile disc florets forming the capitulum Sunflower macro wide.jpg
Inflorescence of sessile disc florets forming the capitulum

Indeterminate or racemose

Indeterminate simple inflorescences are generally called racemose /ˈræsɪms/ . The main kind of racemose inflorescence is the raceme ( /ˈræsm/ , from classical Latin racemus, cluster of grapes). [8] The other kind of racemose inflorescences can all be derived from this one by dilation, compression, swelling or reduction of the different axes. Some passage forms between the obvious ones are commonly admitted.

  • A raceme is an unbranched, indeterminate inflorescence with pedicellate (having short floral stalks) flowers along the axis.
  • A spike is a type of raceme with flowers that do not have a pedicel.
  • A racemose corymb is an unbranched, indeterminate inflorescence that is flat-topped or convex due to their outer pedicels which are progressively longer than inner ones.
  • An umbel is a type of raceme with a short axis and multiple floral pedicels of equal length that appear to arise from a common point. It is characteristic of Umbelliferae.
  • A spadix is a spike of flowers densely arranged around it, enclosed or accompanied by a highly specialised bract called a spathe . It is characteristic of the family Araceae.
  • A flower head or capitulum is a very contracted raceme in which the single sessile flowers share are borne on an enlarged stem. It is characteristic of Dipsacaceae.
  • A catkin or ament is a scaly, generally drooping spike or raceme. Cymose or other complex inflorescences that are superficially similar are also generally called thus.

Determinate or cymose

Determinate simple inflorescences are generally called cymose. The main kind of cymose inflorescence is the cyme (pronounced /sm/ ), from the Latin cyma in the sense 'cabbage sprout', from Greek kuma 'anything swollen'). [9] [10] Cymes are further divided according to this scheme:

  • Only one secondary axis: monochasium
    • Secondary buds always develop on the same side of the stem: helicoid cyme or bostryx
      • The successive pedicels are aligned on the same plane: drepanium
    • Secondary buds develop alternately on the stem : scorpioid cyme
      • The successive pedicels are arranged in a sort of spiral: cincinnus (characteristic of the Boraginaceae and Commelinaceae)
      • The successive pedicels follow a zig-zag path on the same plane: rhipidium (many Iridaceae)
  • Two secondary axes: dichasial cyme
    • Secondary axis still dichasial: dichasium (characteristic of Caryophyllaceae)
    • Secondary axis monochasia: double scorpioid cyme or double helicoid cyme
  • More than two secondary axes: pleiochasium

A cyme can also be so compressed that it looks like an umbel. Strictly speaking this kind of inflorescence could be called umbelliform cyme, although it is normally called simply 'umbel'.

Another kind of definite simple inflorescence is the raceme-like cyme or botryoid; that is as a raceme with a terminal flower and is usually improperly called 'raceme'.

A reduced raceme or cyme that grows in the axil of a bract is called a fascicle. A verticillaster is a fascicle with the structure of a dichasium; it is common among the Lamiaceae. Many verticillasters with reduced bracts can form a spicate (spike-like) inflorescence that is commonly called a spike.

Compound inflorescences

Simple inflorescences are the basis for compound inflorescences or synflorescences. The single flowers are there replaced by a simple inflorescence, which can be both a racemose or a cymose one. Compound inflorescences are composed of branched stems and can involve complicated arrangements that are difficult to trace back to the main branch.

A kind of compound inflorescence is the double inflorescence, in which the basic structure is repeated in the place of single florets. For example, a double raceme is a raceme in which the single flowers are replaced by other simple racemes; the same structure can be repeated to form triple or more complex structures.

Compound raceme inflorescences can either end with a final raceme (homoeothetic), or not (heterothetic). A compound raceme is often called a panicle. This definition is very different from that given by Weberling.

Compound umbels are umbels in which the single flowers are replaced by many smaller umbels called umbellets. The stem attaching the side umbellets to the main stem is called a ray.

The most common kind of definite compound inflorescence is the panicle (of Webeling, or 'panicle-like cyme'). A panicle is a definite inflorescence that is increasingly more strongly and irregularly branched from the top to the bottom and where each branching has a terminal flower.

The so-called cymose corymb is similar to a racemose corymb but has a panicle-like structure. Another type of panicle is the anthela. An anthela is a cymose corymb with the lateral flowers higher than the central ones.

A raceme in which the single flowers are replaced by cymes is called a (indefinite) thyrse. The secondary cymes can be of any of the different types of dichasia and monochasia. A botryoid in which the single flowers are replaced by cymes is a definite thyrse or thyrsoid. Thyrses are often confusingly called panicles.

Other combinations are possible. For example, heads or umbels may be arranged in a corymb or a panicle.

Other

The family Asteraceae is characterised by a highly specialised head technically called a calathid (but usually referred to as 'capitulum' or 'head'). The family Poaceae has a peculiar inflorescence of small spikes ( spikelets ) organised in panicles or spikes that are usually simply and improperly referred to as spike and panicle. The genus Ficus (Moraceae) has an inflorescence called syconium and the genus Euphorbia has cyathia (sing. cyathium), usually organised in umbels.

Some species have inflorescences reduced to composite flowers or pseudanthia, in which case it is difficult to differentiate between inflorescences and single flowers. [11]

Development and patterning

Development

Genetic basis

Genes that shape inflorescence development have been studied at great length in Arabidopsis . LEAFY (LFY) is a gene that promotes floral meristem identity, regulating inflorescence development in Arabidopsis. [12] Any alterations in timing of LFY expression can cause formation of different inflorescences in the plant. [13] Genes similar in function to LFY include APETALA1 (AP1). Mutations in LFY, AP1, and similar promoting genes can cause conversion of flowers into shoots. [12] In contrast to LEAFY, genes like terminal flower (TFL) support the activity of an inhibitor that prevents flowers from growing on the inflorescence apex (flower primordium initiation), maintaining inflorescence meristem identity. [14] Both types of genes help shape flower development in accordance with the ABC model of flower development. Studies have been recently conducted or are ongoing for homologs of these genes in other flower species.

Environmental influences

Inflorescence-feeding insect herbivores shape inflorescences by reducing lifetime fitness (how much flowering occurs), seed production by the inflorescences, and plant density, among other traits. [15] In the absence of these herbivores, inflorescences usually produce more flower heads and seeds. [15] Temperature can also variably shape inflorescence development. High temperatures can impair the proper development of flower buds or delay bud development in certain species, while in others an increase in temperature can hasten inflorescence development. [16] [17] [18]

Meristems and inflorescence architecture

The shift from the vegetative to reproductive phase of a flower involves the development of an inflorescence meristem that generates floral meristems. [19] Plant inflorescence architecture depends on which meristems becomes flowers and which become shoots. [20] Consequently, genes that regulate floral meristem identity play major roles in determining inflorescence architecture because their expression domain will direct where the plant's flowers are formed. [19]

On a larger scale, inflorescence architecture affects quality and quantity of offspring from selfing and outcrossing, as the architecture can influence pollination success. For example, Asclepias inflorescences have been shown to have an upper size limit, shaped by self-pollination levels due to crosses between inflorescences on the same plant or between flowers on the same inflorescence. [21] In Aesculus sylvatica , it has been shown that the most common inflorescence sizes are correlated with the highest fruit production as well. [22]

Related Research Articles

<span class="mw-page-title-main">Panicle</span> Term used in botany to describe a branching of flower heads

A panicle is a much-branched inflorescence. Some authors distinguish it from a compound spike inflorescence, by requiring that the flowers be pedicellate. The branches of a panicle are often racemes. A panicle may have determinate or indeterminate growth.

<span class="mw-page-title-main">Meristem</span> Type of plant tissue involved in cell proliferation

The meristem is a type of tissue found in plants. It consists of undifferentiated cells capable of cell division. Cells in the meristem can develop into all the other tissues and organs that occur in plants. These cells continue to divide until they become differentiated and lose the ability to divide.

<span class="mw-page-title-main">Raceme</span> Unbranched, indeterminate type of inflorescence bearing pedicellate flowers along its axis

A raceme 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 shoot grows in height, with no predetermined growth limit. Examples of racemes occur on mustard, radish, and orchid plants.

<span class="mw-page-title-main">Bract</span> Modified or specialized leaf

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 of a different size, color, shape, or texture. Typically, they also look different from the parts of the flower, such as the petals or sepals. A plant having bracts is referred to as bracteate or bracteolate, while one that lacks them is referred to as ebracteate and ebracteolate, without bracts.

<i>Commelina</i> Genus of flowering plants

Commelina is a genus of approximately 170 species commonly called dayflowers due to the short lives of their flowers. They are less often known as widow's tears. It is by far the largest genus of its family, Commelinaceae. The Swedish taxonomist Carl Linnaeus of the 18th century named the genus after the two Dutch botanists Jan Commelijn and his nephew Caspar, each representing one of the showy petals of Commelina communis.

<span class="mw-page-title-main">Sympodial branching</span>

Sympodial growth is a bifurcating branching pattern where one branch develops more strongly than the other, resulting in the stronger branches forming the primary shoot and the weaker branches appearing laterally. A sympodium, also referred to as a sympode or pseudaxis, is the primary shoot, comprising the stronger branches, formed during sympodial growth. The pattern is similar to dichotomous branching; it is characterized by branching along stems or hyphae.

<i>Aralia spinosa</i> Species of tree

Aralia spinosa, commonly known as devil's walking stick, is a woody species of plant in the genus Aralia of the family Araliaceae. It is native to eastern North America. The various names refer to the viciously sharp, spiny stems, petioles and even leaf midribs. It has also been known as Angelica-tree.

<span class="mw-page-title-main">Corymb</span> Botany term describing a certain type of flower growth

Corymb is a botanical term for an inflorescence with the flowers growing in such a fashion that the outermost are borne on longer pedicels than the inner, bringing all flowers up to a common level. A corymb has a flattish top with a superficial resemblance towards an umbel, and may have a branching structure similar to a panicle. Flowers in a corymb structure can either be parallel, or alternate, and form in either a convex, or flat form.

In botany, an umbel is an inflorescence that consists of a number of short flower stalks that spread from a common point, somewhat like umbrella ribs. The word was coined in botanical usage in the 1590s, from Latin umbella "parasol, sunshade". The arrangement can vary from being flat-topped to almost spherical. Umbels can be simple or compound. The secondary umbels of compound umbels are known as umbellules or umbellets. A small umbel is called an umbellule. The arrangement of the inflorescence in umbels is referred to as umbellate, or occasionally subumbellate.

<span class="mw-page-title-main">Indeterminate growth</span>

In biology and botany, indeterminate growth is growth that is not terminated in contrast to determinate growth that stops once a genetically pre-determined structure has completely formed. Thus, a plant that grows and produces flowers and fruit until killed by frost or some other external factor is called indeterminate. For example, the term is applied to tomato varieties that grow in a rather gangly fashion, producing fruit throughout the growing season. In contrast, a determinate tomato plant grows in a more bushy shape and is most productive for a single, larger harvest, then either tapers off with minimal new growth or fruit or dies.

<span class="mw-page-title-main">Pseudanthium</span> Type of inflorescence, clusters of flowers

A pseudanthium is an inflorescence that resembles a flower. The word is sometimes used for other structures that are neither a true flower nor a true inflorescence. Examples of pseudanthia include flower heads, composite flowers, or capitula, which are special types of inflorescences in which anything from a small cluster to hundreds or sometimes thousands of flowers are grouped together to form a single flower-like structure. Pseudanthia take various forms. The real flowers are generally small and often greatly reduced, but the pseudanthium itself can sometimes be quite large.

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.

<i>Polyscias aemiliguineae</i> Species of shrub

Polyscias aemiliguineae is a species of plant in the family Araliaceae. It is endemic to Réunion.

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>Corymbium</i> Genus of perennial plants from South Africa

Corymbium is a genus of flowering plants in the daisy family comprising nine species. It is the only genus in the subfamily Corymbioideae and the tribe Corymbieae. The species have leaves with parallel veins, strongly reminiscent of monocots, in a rosette and compounded inflorescences may be compact or loosely composed racemes, panicles or corymbs. Remarkable for species in the daisy family, each flower head contains just one, bisexual, mauve, pink or white disc floret within a sheath consisting of just two large involucral bracts. The species are all endemic to the Cape Floristic Region of South Africa, where they are known as plampers.

<i>Adenodaphne</i> Genus of shrubs

Adenodaphne is a genus of shrubs and small trees endemic to New Caledonia belonging to the family Lauraceae. The genus is related to Litsea. They have 12 chromosomes.

<i>Eucomis bicolor</i> Species of flowering plant

Eucomis bicolor, the variegated pineapple lily or just pineapple lily, is a bulbous species of flowering plant in the family Asparagaceae, subfamily Scilloideae, native to Southern Africa. The pale green, purple-margined flowers are arranged in a spike (raceme), topped by a "head" of green leaflike bracts. It is cultivated as an ornamental bulbous plant, although its flowers have an unpleasant smell, attractive to the main pollinators, flies.

<i>Hypericum majus</i> Species of flowering plant

Hypericum majus, the greater Canadian St. John's wort, is a perennial herb native to North America. The specific epithet majus means "larger". The plant has a diploid number of 16.

<i>Cecarria</i> Genus of mistletoes

Cecarria is a monotypic genus in the family Loranthaceae. The sole species is Cecarria obtusifolia, a hemiparasitic aerial shrub.

<i>Scaevola parvifolia</i> Species of plant

Scaevola parvifolia is an erect, many stemmed perennial in the family Goodeniaceae, which is native to Western Australia, the Northern Territory, Queensland and South Australia. It grows to a height of 0.6 m, and its blue-purple flowers may be seen from March to October.

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