Epithallus#Structure

It is defined as the cells above the intercalary meristem; these are not involved in photosynthesis. In Phymatolithon, the epithallium is usually one cell thick, whereas in other genera, such as Pseudolithophyllum, multiple cells exist, with the thickness determined by the difference between their rate of production at the intercalary meristem, and the rate of shedding at the surface;{{Cite journal | last1 = Adey | first1 = W. H. | doi = 10.1007/BF00042707 | title = The genus Pseudolithophyllum (Corallinaceae) in the Gulf of Maine | journal = Hydrobiologia | volume = 27 | issue = 3–4 | pages = 479–497 | year = 1966 | s2cid = 31438562 }} thicknesses of 16 cells or more, spanning 100 μm, have been measured in a representative coralline (Clathromorphum).{{Cite journal | last1 = Adey | first1 = W. H. | doi = 10.1007/BF00045545 | title = The genus Clathromorphum (Corallinaceae) in the Gulf of Maine | journal = Hydrobiologia | volume = 26 | issue = 3–4 | pages = 539–573 | year = 1965 | s2cid = 30943569 }} The thickness is variable within species; in Lithothamnion, a single cell thickness is the norm, but three- or four-cell thick regions are also common.{{Cite journal | last1 = Adey | first1 = W. H. | doi = 10.1007/BF00130389 | title = The genera Lithothamnium, Leptophytum (nov. gen.) and Phymatolithon in the Gulf of Maine | journal = Hydrobiologia | volume = 28 | issue = 3–4 | pages = 321–370 | year = 1966 | s2cid = 2694963 }} The epithallus sometimes overlies the roof of conceptacles, which are exposed only when the overlying epithallus is eventually shed.

The epithallium is less strongly calcified than the underlying cells, facilitating its removal. The meristem itself is the least calcified portion; sometimes there is no mineralization at all, which makes it a plane of weakness where breaking often occurs.

Periodic sloughing of this surface is thought to reduce colonization of corallines by kelp (such as Laminaria),{{Cite journal | last1 = Masaki | first1 = T. | last2 = Fujita | first2 = D. | last3 = Hagen | first3 = N. T. | title = The surface ultrastructure and epithallium shedding of crustose coralline algae in an 'Isoyake' area of southwestern Hokkaido, Japan | doi = 10.1007/BF00027669 | journal = Hydrobiologia | volume = 116 | pages = 218–223 | year = 1984 | s2cid = 40945335 }} epiphytes,{{cite journal | last = Adey | first = W. H. | year = 1964 | title = The genus phymatolithon in the Gulf of Maine | journal = Hydrobiologia | volume = 24 | issue = 1–2 | pages = 377–420 | doi = 10.1007/BF00170412| url = https://deepblue.lib.umich.edu/bitstream/2027.42/42883/1/10750_2004_Article_BF00170412.pdf | hdl = 2027.42/42883 | s2cid = 13005299 | hdl-access = free }}Adey, 1973; Johansen, 1981; Littler & Littler, 1984 and sessile invertebrates.Padilla, 1981{{Cite journal | doi = 10.2307/1938321 | jstor = 1938321 | last1 = Breitburg | first1 = D. L. | title = Residual effects of grazing: inhibition of competitor recruitment by encrusting coralline algae | journal = Ecology | volume = 65 | issue = 4 | pages = 1136–1143 | year = 1984 }} Epithallial cells are covered (in patches) by a cuticle. The deterioration of the outer cells is accelerated in the presence of bacteria.{{Cite journal | last1 = Millson | first1 = C. | last2 = Moss | first2 = B. L. | doi = 10.1515/botm.1985.28.3.123 | title = Ultrastructure of the vegetative thallus of Phymatolithon lenormandii (Aresch. in J. Ag.) Adey | journal = Botanica Marina | volume = 28 | issue = 3 | pages = 123 | year = 1985 | s2cid = 84471882 }}

A similar mechanism is found in geniculate reds.(Borowitzka & Vesk, 1978) Epidermal tissue is also shed by unrelated algae: the fleshy reds and browns,{{cite journal |author=Filion-Myklebust, CC.|author2=T.A. Norton |year=1981 |title=Epidermis shedding in the brown seaweed Ascophyllum nodosum (L.) Le Jolis |journal=Marine Biology Letters | pages= 45–51 |volume= 2}} (e.g. Chondrus, Ascophyllum;{{Cite journal|author1-link=John McNeill Sieburth | last1 = Sieburth | first1 = J. M. | last2 = Tootle | first2 = J. L. | doi = 10.1111/j.1529-8817.1981.tb00819.x | title = Seasonality of microbial fouling on Ascophyllum nodosum (L.) Lejol., Fucus vesiculosus L., Polysiphonia lanosa (L.) Tandy and Chondrus crispus Stackh | journal = Journal of Phycology | volume = 17 | pages = 57–64 | year = 1981 | s2cid = 86258108 }} Halidrys,{{cite journal | title= The control of epiphytes by Halidrys siliquosa (L.) Lyngb. (Phaeophyta, Cytoseiraceae) | author = Moss, B L| journal=Phycologia |volume= 21| issue= 2| pages= 185–188|year= 1982 | doi= 10.2216/i0031-8884-21-2-185.1}} Himanthalia{{cite journal

| title = Epiphyte survival on skin-shedding macrophytes

| author1= Russell, G |author2= Veltkamp, CJ

| journal=Marine Ecology Progress Series

| volume= 18

| issue= 1–2

| pages= 149–153

| year = 1984

| doi = 10.3354/meps018149

| bibcode= 1984MEPS...18..149R | doi-access= free

}}) and the calcaerous greens.{{Cite journal | last1 = Borowitzka | first1 = M. A. | last2 = Larkum | first2 = A. W. D. | doi = 10.1111/j.1529-8817.1977.tb02879.x | title = Calcification in the green alga Halimeda. I. An ultrastructure study of thallus development | journal = Journal of Phycology | volume = 13 | pages = 6–16 | year = 1977 | s2cid = 86035589 }} (cited to support this fact in Johnson & Mann 1989, although I missed how the paper supports it) Some sea grasses also periodically shed their external cell walls to avoid epiphyte cover.{{Cite journal | last1 = Jagels | first1 = R. | title = Studies of a marine grass, Thalassia testudinum. I. Ultrastructure of the osmoregulatory leaf cells | doi = 10.2307/2441514 | journal = American Journal of Botany | volume = 60 | issue = 10 | pages = 1003–1009 | year = 1973 | jstor = 2441514 }}; also {{cite journal|author=Jagels, R. H.|year= 1970 |title=Cell wall development in a marine monocotyledon|journal=American Journal of Botany |volume=57|issue= 6 |pages=737–738|jstor=2441153}} In the browns, this is accomplished by shedding cell wall material, without damaging the underlying cells.

The epithallus probably originated from cover cells, which are considered to be homologous structures.

For a cross-sectional image in Clathromorphum circumscriptum, see plate 38 (p. 415) in Adey, 1964 (referenced below)

Additional images showing the epithallus can be seen in Masaki et al. (1984).

NB incomplete citations refer to references in Johnson & Mann (1986).

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Category:Red algae