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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

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]

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

Indeterminate or racemose

Indeterminate simple inflorescences are generally called racemose /ˈræsɪmoʊs/. The main kind of racemose inflorescence is the raceme (/ˈræsiːm/, 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.

Determinate or cymose

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

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]

References

  1. ^ Guertin, P., Barnett, L., Denny, E.G., Schaffer, S.N. 2015. USA National Phenology Network Botany Primer. USA-NPN Education and Engagement Series 2015-001. www.usanpn.org.
  2. ^ "Inflorescence | Racemes, Spikes & Cymes | Britannica". www.britannica.com. 2023-09-26. Retrieved 2023-11-03.
  3. ^ Kirchoff, Bruce K.; Claßen-Bockhoff, Regine (2013). "Inflorescences: concepts, function, development and evolution". Annals of Botany. 112 (8): 1471–6. doi:10.1093/aob/mct267. PMC 3828949. PMID 24383103.
  4. ^ "PlantNET - NSW Flora Online - Glossary". Retrieved May 17, 2022.
  5. ^ Fernanda Martínez-Velarde, Maria; 6 others, and (2023). "Desmopsisterriflora, an extraordinary new species of Annonaceae with flagelliflory". PhytoKeys (227): 181–198. doi:10.3897/phytokeys.227.102279. PMC 10314296. PMID 37396012.((cite journal)): CS1 maint: numeric names: authors list (link)
  6. ^ Bradley, Desmond; Ratcliffe, Oliver; Vincent, Coral; Carpenter, Rosemary; Coen, Enrico (1997-01-03). "Inflorescence Commitment and Architecture in Arabidopsis". Science. 275 (5296): 80–83. doi:10.1126/science.275.5296.80. ISSN 0036-8075. PMID 8974397. S2CID 20301629.
  7. ^ Kubitzki, Klaus, and Clemens Bayer. 2002. Flowering plants, Dicotyledons: Malvales, Capparales, and non-betalain Caryophyllales. The Families and genera of vascular plants, 5. Berlin: Springer. p. 77
  8. ^ Oxford English Dictionary. Raceme 2. Bot. A type of inflorescence in which the flowers are arranged on short, nearly equal, lateral pedicels, at equal distances along a single elongated axis
  9. ^ Collins English Dictionary. 8th Edition first published in 2006
  10. ^ Oxford English Dictionary. Cyme(1) Bot. A species of inflorescence wherein the primary axis bears a single terminal flower which develops first, the system being continued by axes of secondary and higher orders which develop successively in like manner; a centrifugal or definite inflorescence: opposed to Raceme. Applied esp. to compound inflorescences of this type forming a more or less flat head.
  11. ^ Tucker, Shirley C.; Grimes, James (1999-10-01). "The inflorescence: Introduction". The Botanical Review. 65 (4): 303–316. Bibcode:1999BotRv..65..303T. doi:10.1007/BF02857752. ISSN 0006-8101. S2CID 29599096.
  12. ^ a b Shannon, S.; Meeks-Wagner, D. R. (1993-06-01). "Genetic Interactions That Regulate Inflorescence Development in Arabidopsis". The Plant Cell. 5 (6): 639–655. doi:10.1105/tpc.5.6.639. ISSN 1040-4651. PMC 160302. PMID 12271079.
  13. ^ Schultz, E. A.; Haughn, G. W. (1991-08-01). "LEAFY, a Homeotic Gene That Regulates Inflorescence Development in Arabidopsis". The Plant Cell. 3 (8): 771–781. doi:10.1105/tpc.3.8.771. ISSN 1040-4651. PMC 160044. PMID 12324613.
  14. ^ Alvarez, John; Guli, Catherine L.; Yu, Xiang-Hua; Smyth, David R. (1992-01-01). "terminal flower: a gene affecting inflorescence development in Arabidopsis thaliana". The Plant Journal. 2 (1): 103–116. doi:10.1111/j.1365-313X.1992.00103.x. ISSN 1365-313X.
  15. ^ a b Louda, Svata M.; Potvin, Martha A. (1995-01-01). "Effect of Inflorescence-Feeding Insects on the Demography and Lifetime of a Native Plant". Ecology. 76 (1): 229–245. Bibcode:1995Ecol...76..229L. doi:10.2307/1940645. ISSN 1939-9170. JSTOR 1940645.
  16. ^ Moss, G (27 November 2015). "Influence of Temperature and Photoperiod on Flower Induction and Inflorescence Development in Sweet Orange (Citrus Sinensis L. Osbeck)". Journal of Horticultural Sciences. 44 (4): 311–320. doi:10.1080/00221589.1969.11514314.
  17. ^ Bjorkman, T.; Pearson, K. J. (1998-01-01). "High temperature arrest of inflorescence development in broccoli (Brassica oleracea var. italica L.)". Journal of Experimental Botany. 49 (318): 101–106. doi:10.1093/jxb/49.318.101. ISSN 0022-0957.
  18. ^ BREWSTER, J. L. (1983-04-01). "Effects of Photoperiod, Nitrogen Nutrition and Temperature on Inflorescence Initiation and Development in Onion (Allium cepa L.)". Annals of Botany. 51 (4): 429–440. doi:10.1093/oxfordjournals.aob.a086487. ISSN 0305-7364.
  19. ^ a b Souer, E.; Krol, A. van der; Kloos, D.; Spelt, C.; Bliek, M.; Mol, J.; Koes, R. (1998-02-15). "Genetic control of branching pattern and floral identity during Petunia inflorescence development". Development. 125 (4): 733–742. doi:10.1242/dev.125.4.733. ISSN 0950-1991. PMID 9435293.
  20. ^ Benlloch, R.; Berbel, A.; Serrano-Mislata, A.; Madueno, F. (2007-09-01). "Floral Initiation and Inflorescence Architecture: A Comparative View". Annals of Botany. 100 (3): 659–676. doi:10.1093/aob/mcm146. ISSN 0305-7364. PMC 2759223. PMID 17679690.
  21. ^ WYATT, ROBERT (1980-05-01). "The Reproductive Biology of Asclepias Tuberosa: I. Flower Number, Arrangement, and Fruit-Set". New Phytologist. 85 (1): 119–131. doi:10.1111/j.1469-8137.1980.tb04453.x. ISSN 1469-8137.
  22. ^ Wyatt, Robert (1982-04-01). "Inflorescence Architecture: How Flower Number, Arrangement, and Phenology Affect Pollination and Fruit-Set". American Journal of Botany. 69 (4): 585–594. doi:10.1002/j.1537-2197.1982.tb13295.x. ISSN 1537-2197. JSTOR 2443068.

Bibliography

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