Anthocerotophyta

Hornworts are a group of non-vascular plants constituting the division Anthocerotophyta. The common name refers to the elongated horn-like structure, which is the sporophyte. As in mosses and liverworts, the flattened, green plant body of a hornwort is the gametophyte plant. Hornworts may be found worldwide, though they tend to grow only in places that are damp or humid. Some species grow in large numbers as tiny weeds in the soil of gardens and cultivated fields. Large tropical and sub-tropical species of Dendroceros may be found growing on the bark of trees. The total number of species is still uncertain. While there are more than 300 published species names, the actual number could be as low as 100-150 species. The plant body of a hornwort is a haploid gametophyte stage. This stage usually grows as a thin rosette or ribbon-like thallus between one and five centimeters in diameter. Each cell of the thallus usually contains just one chloroplast. In half of the roughly 200 hornwort species, this chloroplast is fused with other organelles to form a large pyrenoid that both enables more efficient photosynthesis and stores food. The pyrenoid is comprised predominantly of RuBisCO, the key enzyme in carbon fixation. By using inorganic carbon transporters and carbonic anhydrases, up to a 50-fold increase in CO2 levels can be achieved. This particular feature is very unusual in land plants, unique to hornworts, but is common among algae. Many hornworts develop internal mucilage-filled cavities or canals when groups of cells break down. They will secrete hormogonium-inducing factors (HIF) that stimulate nearby, free-living photosynthetic cyanobacteria, especially species of Nostoc, to invade and colonize these cavities. Such colonies of bacteria growing inside the thallus give the hornwort a distinctive blue-green color. There may also be small slime pores on the underside of the thallus. These pores superficially resemble the stomata of other plants. The horn-shaped sporophyte grows from an archegonium embedded deep in the gametophyte. The sporophyte of a hornwort is unusual in that it grows from a meristem near its base, instead of from its tip the way other plants do. Unlike liverworts, most hornworts have true stomata on their sporophyte as mosses do. The exceptions are the genera Notothylas and Megaceros, which do not have stomata. The sporophyte of most hornworts are also photosynthetic, which is not the case with liverworts. When the sporophyte is mature, it has a multicellular outer layer, a central rod-like columella running up the center, and a layer of tissue in between that produces spores and pseudo-elaters. The pseudo-elaters are multi-cellular, unlike the elaters of liverworts. They have helical thickenings that change shape in response to drying out; they twist and thereby help to disperse the spores. Hornwort spores are relatively large for bryophytes, measuring between 30 and 80 µm in diameter or more. The spores are polar, usually with a distinctive Y-shaped tri-radiate ridge on the proximal surface, and with a distal surface ornamented with bumps or spines The life of a hornwort starts from a haploid spore. In most species, there is a single cell inside the spore, and a slender extension of this cell called the germ tube germinates from the proximal side of the spore. The tip of the germ tube divides to form an octant (solid geometry) of cells, and the first rhizoid grows as an extension of the original germ cell. The tip continues to divide new cells, which produces a thalloid protonema. By contrast, species of the family Dendrocerotaceae may begin dividing within the spore, becoming multicellular and even photosynthetic before the spore germinates. In either case, the protonema is a transitory stage in the life of a hornwort.