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 formed. 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.
One can also define an inflorescence as the reproductive portion of a plant that bears a cluster of flowers in a specific pattern.
The stem holding the whole inflorescence is called a peduncle and the major axis holding the flowers or more branches within the inflorescence is called the rachis. The stalk of each single flower 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 or complex. The rachis may be one of several types, including single, composite, umbel, spike or raceme.
General characteristicsInflorescences 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.
BractsInflorescences usually have modified shoots 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 main stem 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 can distinguish:
- Ebracteate inflorescences: No bracts in the inflorescence.
- Bracteate inflorescences: The bracts in the inflorescence are very specialised, sometimes reduced to small scales, divided or dissected.
- Leafy inflorescences: Though often reduced in size, the bracts are unspecialised and look like the typical leaves of the plant, so that the term flowering stem is usually applied instead of inflorescence. This use is not technically correct, as, despite their 'normal' appearance, these leaves are considered, in fact, bracts, so that 'leafy inflorescence' is preferable.
- Leafy-bracted inflorescences: Intermediate between bracteate and leafy inflorescence.
Terminal flowerPlant 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 inflorescence: Monopodial growth. The terminal bud keeps growing and forming lateral flowers. A terminal flower is never formed.
- Determinate inflorescence: Sympodial growth. The terminal bud forms a terminal flower and then dies out. Other flowers then grow from lateral buds.
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 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, while the others tend to mature starting from the bottom of the stem. This pattern is called acropetal maturation. When flowers start to mature from the top of the stem, maturation is basipetal, while when the central mature first, divergent.
PhyllotaxisAs 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 is located at the axil of a bract, the location of the bract in relation to the stem holding the flower is indicated by the use of different terms and may be a useful diagnostic indicator.
Typical placement of bracts include:
- Some plants have bracts that subtend the inflorescence, where the flowers are on branched stalks; the bracts are not connected to the stalks holding the flowers, but are adnate or attached to the main stem
- Other plants have the bracts subtend the pedicel or peduncle of single flowers.
- When the bract is attached to the stem holding the flower, it is said to be recaulescent; sometimes these bracts or bracteoles are highly modified and appear to be appendages of the flower calyx. Recaulescences is the fusion of the subtending leaf with the stem holding the bud or the bud itself, thus the leaf or bract is adnate to the stem of flower.
- When the formation of the bud is shifted up the stem distinctly above the subtending leaf, it is described as concaulescent.
Indeterminate or racemoseIndeterminate simple inflorescences are generally called racemose. The main kind of racemose inflorescence is the raceme. 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 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
- 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 : scorpoid cyme
- ** The successive pedicels are arranged in a sort of spiral: cincinnus
- ** The successive pedicels follow a zig-zag path on the same plane: rhipidium
- Two secondary axes: dichasial cyme
- * Secondary axis still dichasial: dichasium
- * Secondary axis monochasia: double scorpioid cyme or double helicoid cyme
- More than two secondary axes: pleiochasium
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 inflorescence that is commonly called a spike.
Compound inflorescencesSimple 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, or not. A compound raceme is often called a panicle. Note that 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. 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 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.
OtherThe family Asteraceae is characterised by a highly specialised head technically called a calathid. The family Poaceae has a peculiar inflorescence of small spikes organised in panicles or spikes that are usually simply and improperly referred to as spike and panicle. The genus Ficus has an inflorescence called syconium and the genus Euphorbia has cyathia, usually organised in umbels.
Development and patterning
Genetic basisGenes that shape inflorescence development have been studied at great length in Arabidopsis. LEAFY is a gene that promotes floral meristem identity, regulating inflorescence development in Arabidopsis. Any alterations in timing of LFY expression can cause formation of different inflorescences in the plant. Genes similar in function to LFY include APETALA1. Mutations in LFY, AP1, and similar promoting genes can cause conversion of flowers into shoots. In contrast to LEAFY, genes like terminal flower support the activity of an inhibitor that prevents flowers from growing on the inflorescence apex, maintaining inflorescence meristem identity. 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 influencesInflorescence-feeding insect herbivores shape inflorescences by reducing lifetime fitness, seed production by the inflorescences, and plant density, among other traits. In the absence of this herbivory, inflorescences usually produce more flower heads and seeds. 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.
Meristems and inflorescence architectureThe shift from the vegetative to reproductive phase of a flower involves the development of an inflorescence meristem that generates floral meristems. Plant inflorescence architecture depends on which meristems becomes flowers and which become shoots. 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.
On a larger scale, inflorescence architecture affects the 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. In Aesculus sylvatica, it has been shown that the most common inflorescence sizes are correlated with the highest fruit production as well.