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Lichen morphology describes the external appearance and structures of a lichen. These complex, symbiotic organisms are composed of a fungus (the mycobiont) in association with an alga and/or a cyanobacteria (the photobiont). Morphology can vary considerably from lichen to lichen.
Overview
[edit]Lichens are composite organisms made up of multiple species. A fungal partner (known as the mycobiont), combines with one or more photosynthetic partners (known as photobionts); sometimes a yeast is also involved.[1] Each partner contributes to the organism's success. In most lichens, the fungal partner provides the structure in which the various partners live; this structure helps to protect the photobiont from the outside environment.[2] The photosynthetic partner(s) provide the nutrients which the various partners need to survive.[2] The yeast (where present) appears to help ward off microbes and potential predators through the production of various chemicals.[1] Thallus types have evolved to provide the lichen's photobiont with optimal levels of light, water, and carbon dioxide, with different environmental conditions favouring different forms.[3][4]
Thallus
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The thallus (pl. thalli) makes up the bulk of most lichens. This vegetative tissue provides the lichen's overall structure, and houses its various partners. The thallus's growth form depends on the species involved. Crustose lichens are crust-like, as their name suggests. They are tightly bound to the substrate upon which they grow (looking like a paint splotch) and cannot be removed without at least partial destruction of either the lichen or the substrate. Foliose lichens are flattened and leaf-like, and can be at least partially lifted from the substrate on which they grow. Fruticose lichens are either threadlike or shrubby and coral-like, and are only minimally attached to the substrate on which they grow. Squamulose lichens are scale-like, typically attached only along one edge and often overlapping, like shingles on a roof. The thallus may or may not be covered with a cortex, and may or may not give rise to various vegetative and reproductive structures.
In most lichens, the fungal partner makes up the bulk of the thallus and determines its shape, though in a few cases, the photosynthetic partner instead creates the structure. Regardless of which partner is "in charge", an association between the partners (and in some cases, a yeast) is required for the development of the lichen's typical growth form. When grown in the laboratory in the absence of its photobiont partner(s), a lichen fungus develops as an undifferentiated mass of hyphae. Only when combined with its photobiont under appropriate conditions does its characteristic form emerge, in the process called morphogenesis. In a few remarkable cases, a single lichen fungus can develop into very different forms depending on whether it associates with a green alga or a cyanobacteria. These alternative forms were originally thought to be different species, until they were found growing together in a conjoined manner.
Cortex
[edit]Many lichens have a cortex on one or both of their surfaces (upper and lower). This outermost "skin", which is composed of tightly woven fungal hyphae arranged in a highly uniform manner, can be up to several hundred micrometers thick (less than a millimeter / less than a hundredth of an inch). It helps to protect the lichen from other organisms and from various environmental factors – especially desiccation and excessive ultraviolet light. Powdery lichens like those in the genus Lepraria and wispy lichens like those in the genus Coenogonium lack any cortex at all. Those crustose lichens which have a cortex have one only on their upper surface, while most foliose lichens have a separate cortex on their upper and lower surfaces. Fruticose lichens have a single cortex which extends all the way around their surface.
Secondary metabolites produced by the lichen are typically stored in the cortex. These may include pigments which help to cut the amount of UV light which reaches the photobiont, or compounds which make the lichen unpalatable to other organisms.
Vegetative structures
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Cephalodium
[edit](pl. cephalodia) Lichens which contain both algal and cyanobacterial photobionts house the latter in small, gall-like structures known as cephalodia. These may be on the upper or lower surface of the thallus or within its interior. While most are simply shaped – typically round, sac-like, or lobed – a few are more complexly branched and coral-like.[5] They range in size from "minute" to several millimeters across and are typically a different color than the rest of the lichen's thallus.[6][5]
Cilium
[edit](pl. cilia) Cilia are fine, hair-like outgrowths of either the thallus or the apothecium. They are common in many foliose and some fruticose lichens.[7]
Cyphella
[edit](pl. cyphellae) A sharply defined, rounded, ovate, or shapeless pore in the lower thallus surface (typically the lower cortex), which is lined with a "pseudocortex" made of loosely connected, non-gelatinized hyphae (often with globular cells, formed from the medulla) and bounded by a pale ring; known to occur in the genera Sticta and Oropogon.[8]
Holdfast
[edit]A holdfast is hyphae
Podetium
[edit](pl. podetia) Podetia are hollow, upright, secondary thalli which rise from the lichen's primary thallus in a small number of species – primarily those of the genus Cladonia. Podetia may be spike-like, branched, or cup-shaped. Where they occur, they support the lichen's reproductive structures. They are formed of "generative" tissue.
Pseudopodetium
[edit](pl. pseudopodetia) Solid, upright stalks originating from the thallus. They are similar to podetia, but are made of vegetative (rather than generative) tissue.[9] They are associated with the genera Baeomyces, Dibaeis, Leprocaulon, Pilophorus, and Stereocaulon.[10]
Rhizine
[edit]Foliose lichens often have small, root-like strands, known as rhizines, extending from their lower cortex. These help to attach the lichen to its substrate, but do not transport any water or nutrients. They come in various shapes; some are simple threads, others are forked or complexly branched. The density and shape of its rhizines can aid in a lichen's identification.
Reproductive structures
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The vast majority of lichen species reproduce sexually. They use specialized structures to do so.[citation needed] A smaller number of species reproduce asexually as well. These too use one or more of several specialized structures.[citation needed]
Apothecium
[edit]An apothecium (pl. apothecia) is a wide, open, saucer-shaped or cup-shaped fruit body. It is sessile and fleshy. The structure of the apothecium chiefly consists of three parts: hymenium (upper concave surface), hypothecium, and excipulum (the "foot"). The asci are present in the hymenium layer. The asci are freely exposed at maturity.
Ascocarp
[edit]The fruiting body of an ascomycete fungus, containing the asci and ascospores.
Blastidium
[edit](pl. blastidia) Blastidia are tiny, rounded, granule-like propagules budding; subsequent blastidia are formed from the tips of the previous ones.[11][7]
Isidium
[edit](pl. isidia) A propagule that is an outgrowth of the thallus; it has a cortex and contains photobiont cells.
Perithecium
[edit]Plural perithecia. A spherical or flask-shaped ascocarp that is sessile or partly immersed in the thallus, with a single opening (ostiole) and enclosed by a distinct wall; a characteristic of pyrenolichens.[12] Although it was in 1831 that Elias Fries first applied the term perithecium to lichen fruit bodies, the word was originally coined by Christiaan Hendrik Persoon in 1794.[13]
Pycnidium
[edit](pl. pycnidia) An asexual fruiting body, or conidium, that is typically round, obpyriform, or lageniform. It has a circular or elongated ostiole that has an inner surface lined with conidiophores.
Soralium
[edit](pl. soralia A part of the thallus where the cortex has cracked or broken down and soredia are produced. Soralia can be further characterized as diffuse if they are spread out on the upper thallus surface as a continuous layer, or delimited if they are confined to a more restricted area. If soralia originate in tubercules they are tuberculate, while they are fissural if they are created in fissures.[14]
Soredium
[edit](pl. soredia) In some lichens, the surface of a thallus may show minute, powdery granules (called soredia), each soredium consisting of a few photobiont cells surrounded by fungal filaments.
See also
[edit]References
[edit]- ^ a b Van Hoose 2021.
- ^ a b Baron 1999, p. 14.
- ^ Büdel & Scheidegger 1996, p. 37.
- ^ Aragón et al. 2019, p. 8.
- ^ a b Honegger 2008, p. 86.
- ^ Budel & Scheidegger 2008, p. 56.
- ^ a b Brodo, Sharnoff & Sharnoff 2001, p. 756.
- ^ Kirk et al. 2008, p. 187.
- ^ Ulloa & Hanlin 2012, p. 517.
- ^ Smith et al. 2009, p. 36.
- ^ Smith et al. 2009, p. 22.
- ^ Ulloa & Hanlin 2012, p. 463.
- ^ Mitchell 2014, p. 17.
- ^ Ulloa & Hanlin 2012, p. 586.
Sources
[edit]- Aragón, Gregorio; Martínez, Isabel; Hurtado, Pilar; Benítez, Ángel; Rodríguez, Clara; Prieto, María (2019). "Using growth forms to predict epiphytic lichen abundance in a wide variety of forest types". Diversity. 11 (4): 51. doi:10.3390/d11040051.
- Baron, George (1999). Understanding Lichens. Slough: Richmond Publishing. ISBN 978-0-85546-252-9.
- Büdel, B.; Scheidegger, C. (1996). "Thallus morphology and anatomy". In Nash, Thomas (ed.). Lichen Biology. Cambridge: Cambridge University Press. ISBN 978-0-521-45368-4.
- Budel, Burkhard; Scheidegger, Christoph (2008). "Thallus morphology and anatomy". In Nash III, Thomas E. (ed.). Lichen Biology (2nd ed.). Cambridge: Cambridge University Press. ISBN 978-0-521-87162-4.
- Honegger, Rosmarie (2008). "Morphogenesis". In Nash III, Thomas E. (ed.). Lichen Biology (2nd ed.). Cambridge: Cambridge University Press. ISBN 978-0-521-87162-4.
- Van Hoose, Natalie (21 July 2021). "Yeast emerges as hidden third partner in lichen symbiosis". Purdue University News. Purdue University.