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10.4: Green Algae- Precursors of Land Plants

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    Streptophytes

    Until recently, all photosynthetic eukaryotes were considered members of the kingdom Plantae. The brown, red, and gold algae, however, have been reassigned to the Protista kingdom. This is because apart from their ability to capture light energy and fix CO2, they lack many structural and biochemical traits that distinguish plants from protists. The position of green algae is more ambiguous. Green algae contain the same carotenoids and chlorophyll a and b as land plants, whereas other algae have different accessory pigments and types of chlorophyll molecules in addition to chlorophyll a. Both green algae and land plants also store carbohydrates as starch. Cells in green algae divide along cell plates called phragmoplasts, and their cell walls are layered in the same manner as the cell walls of embryophytes. Consequently, land plants and closely related green algae are now part of a new monophyletic group called Streptophyta.

    The remaining green algae, which belong to a group called Chlorophyta, include more than 7000 different species that live in fresh or brackish water, in seawater, or in snow patches. A few green algae even survive on soil, provided it is covered by a thin film of moisture in which they can live. Periodic dry spells provide a selective advantage to algae that can survive water stress. Some green algae may already be familiar, in particular Spirogyra and desmids. Their cells contain chloroplasts that display a dizzying variety of shapes, and their cell walls contain cellulose, as do land plants. Some green algae are single cells, such as Chlorella and Chlamydomonas, which adds to the ambiguity of green algae classification, because plants are multicellular. Other algae, like Ulva (commonly called sea lettuce), form colonies (Figure 1).

    Light micrograph A shows rectangular Spirogyra cells linked in a chain. Light micrograph B shows a oval green desmid cell. Electron micrograph C shows egg-shaped Chlamydomonas cells attached to thin stalks. Photo D shows a colony of Ulva that resembles leaf lettuce.
    Figure 1. Chlorophyta include (a) Spirogyra, (b) desmids, (c) Chlamydomonas, and (d) Ulva. Desmids and Chlamydomonas are single-celled organisms, Spirogyra forms chains of cells, and Ulva forms colonies resembling leaves (credit b: modification of work by Derek Keats; credit c: modification of work by Dartmouth Electron Microscope Facility, Dartmouth College; credit d: modification of work by Holger Krisp; scale-bar data from Matt Russell)

    Reproduction of Green Algae

    Green algae reproduce both asexually, by fragmentation or dispersal of spores, or sexually, by producing gametes that fuse during fertilization. In a single-celled organism such as Chlamydomonas, there is no mitosis after fertilization. In the multicellular Ulva, a sporophyte grows by mitosis after fertilization (and thus exhibits alternation of generations). Both Chlamydomonas and Ulva produce flagellated gametes.

    Charales

    Chara is a water plant with a long, flexible green stem. Whorls of slender stalks periodically radiate from the stem.
    Figure 2. The representative alga, Chara, is a noxious weed in Florida, where it clogs waterways. (credit: South Florida Information Access, U.S. Geological Survey)

    The charophytes include several different algal orders that have each been suggested to be the closest relatives of the land plants: the Charales, the Zygnematales, and the Coleochaetales. The Charales can be traced back 420 million years. They live in a range of freshwater habitats and vary in size from a few millimeters to a meter in length. The representative genus is Chara (Figure 25.8), often called muskgrass or skunkweed because of its unpleasant smell. Large cells form the thallus: the main stem of the alga. Branches arising from the nodes are made of smaller cells. Male and female reproductive structures are found on the nodes, and the sperm have flagella. Although Chara looks superficially like some land plants, a major difference is that the stem has no supportive tissue. However, the Charales exhibit a number of traits that are significant for adaptation to land life. They produce the compounds lignin and sporopollenin, and form plasmodesmata that connect the cytoplasm of adjacent cells. Although the life cycle of the Charales is haplontic (the main form is haploid, and diploid zygotes are formed but have a brief existence), the egg, and later, the zygote, form in a protected chamber on the haploid parent plant.

    The Coleochaetes are branched or disclike multicellular forms. They can produce both sexually and asexually, but the life cycle is basically haplontic. Recent extensive DNA sequence analysis of charophytes indicates that the Zygnematales are more closely related to the embryophytes than the Charales or the Coleochaetales. The Zygnematales include the familiar genus Spirogyra, as well as the desmids. As techniques in DNA analysis improve and new information on comparative genomics arises, the phylogenetic connections between the charophytes and the land plants will continued to be examined to produce a satisfactory solution to the mystery of the origin of land plants.

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