Themes > Science > Botanical Sciences > Classification of Plants > Cryptogamia (Seedless Plants) > Bryophyta Mosses and Liverworts > Nonvascular Plants


Nonvascular plants are the simplest of all land dwelling plants.  Like their closest ancestors, the green algae, they lack an internal means for water transportation.  They also do not produce seeds or flowers.  They generally only reach a height of one to two centimeters, because they lack the woody tissue necessary for support on land.
 
Evolutionary History

Scientists possess fossil records that lead them to believe that plants evolved during four distinct periods.  Nonvascular plants arose first, during the late Ordovician period of the Paleozoic Era, approximately 460 million years ago.  Their closest non plant ancestor is a type of green algae called charophytes.  Nonvascular plants exhibit several homologies to charophytes, such as

  • Homologous chloroplasts - Both nonvascular plants and charophytes use chlorophyll and beta carotene as accessory pigments in their chloroplasts.  Also, the thylakoid membranes of both are stacked as grana.
  • Biochemical Similarities - Charophytes and plants both have cellulose in their cell walls, and their peroxisomes have the same enzyme composition.
  • Similarities in the mechanisms of mitosis and cytokinesis - During mitosis in both nonvascular plants and charophytes, the nuclear envelope completely disperses during late prophase.  In cytokinesis, the formation of a cell plate involves the cooperation of microtubules, actin microfilaments, and vesicles.
  • Similarity in sperm ultrastructure - In the details of sperm ultrastructure, charophytes are more similar to plants than to other green algae.
  • Genetic relationship - Nonvascular plant nuclear genes and ribosomal RNA most closely resemble those of charophytes.
Scientists also believe that living in shallow water was a preadaptation to living on land.  Natural selection probably favored algae (living on the fringes of bodies of water) that could survive through periods when they were not submerged.  Waxy cuticles and jacketed organs, both characteristics of nonvascular plants, are possible adaptations that the algae developed to survive in these conditions.

Reproduction

Nonvascular plants can reproduce both sexually and asexually.  Asexual reproduction is the less common method.  It basically consists of the regeneration of plant material, leaves or other parts, that fall to the ground and generate secondary plants which bear new buds.

Most nonvascular plants, though, reproduce sexually.  Their gametes develop within structures called gametangia, which are organs that have protective jackets of sterile cells that prevent the gametes from drying out during development.  The male gametangium is called the antheridium, and it produces flagellated sperm.  The female gametangium, or archegonium, produces a single egg (ovum).  As in algae, the flagellated sperm require water to swim from the antheridium to the archegonium and fertilize the egg.  For most species, a film of rainwater or dew is sufficient for fertilization.
Nonvascular plants, along with all other members of the plant world, engage in a life cycle known as alternation of generations.  During this cycle, the plant forms both a multicellular diploid generation, the sporophyte, and a multicellular haploid generation, the gametophyte.  In nonvascular plants, the dominant generation is the gametophyte, whereas in most plants the sporophyte is dominant. The sporophyte in nonvascular plants is smaller and shorter lived than the gametophyte, and it depends on the gametophyte for survival.
After the sperm swims to the archegonium and fertilizes the egg, the diploid zygote divides by mitosis and develops into an embryonic sporophyte within the archegonium.  This sporophyte grows into a long stalk whose base remains attached to the archegonium as the top emerges.  A sporangium forms at the tip of the stalk, and haploid spores develop within it by meiosis.  Eventually, the sporangium bursts and the spores scatter.  They germinate by mitotic division and form structures known as protonema, which eventually develop into mature gametophytes, completing the life cycle.   




Habitats
    

Nonvascular plants are almost always found in damp, shady places.  They have little or no resistance to drying, and because they lack vascular tissue they cannot carry water from the ground to the aerial parts of the plant.  Like sponges, they must imbibe the water lying on their surfaces and distribute it by the relatively slow means of diffusion, capillary action, and cytoplasmic streaming.  Therefore, they cannot survive for very long in areas that are not constantly moist.  Some can survive in alternative habitats such as sand dunes, but the majority thrive in dark, dank places.  Because of their limited range of terrestrial habitats, nonvascular plants have never dominated much of the earth's landscape.

Energy Acquisition

Like most plants, nonvascular plants acquire energy through photosynthesis.  During this process, the plant converts light energy into chemical energy, then proceeds to store it in the form of glucose or other organic compounds.  Photosynthesis usually occurs in the upper parts of nonvascular plants, where they produce many small stemlike and leaflike appendages.

Divisions

There are three divisions of nonvascular plants: Bryophyta, Hepatophyta, and Anthocerophyta.  Until recently, scientists grouped all three together as one division, Bryophyta, but the current view is that they are probably not related.  They do share some key characteristics, such as the presence of a waxy cuticle and gametangia, but they all have distinct characteristics that warrant separate divisions. 



Bryophyta

Division Bryophyta consists of the mosses, and it includes approximately 10,000 species.  Mosses are the most common and familiar nonvascular plants.  They usually grow in a mat formation, which consists of many plants growing in a tight pack to hold one another up.  The mat usually has a spongy quality which enables it to retain water, thus aiding in reproduction and preventing the plant from drying out.  Mosses possess multicellular, rootlike structures known as rhizoids which they use for attachment and water absorption.  All mosses consist of "stems", either branched or unbranched, that bear leaflike structures.  It is important to note that these "stems", "roots", and "leaves" are not homologous to those of vascular plants.

In mosses, the capsules of the sporophytes (sporangia) are much more complex than those of the other nonvascular plants.  In many cases the sporangia will possess a lid, or operculum, which is separated from the rest of the capsule by a ring.  This lid will eventually detach, releasing spores.  Spore release is sometimes aided by the movement of peristome teeth, which are arranged radially around the mouth of the capsule.

Hepatophyta   

Division Hepatophyta is the home of the liverworts: it contains approximately 6,500 species.  These plants differ from mosses in that many do not have their characteristic stem/leaf structure.  Instead, their bodies are divided into deeply grooved lobes.  Some have coil shaped cells in their sporangia which spring out of the capsule when it opens, helping to disperse the spores.  Their capsules are usually much simpler than those of the mosses, consisting of simple spheres that split longitudinally into four sections when mature.  Some also develop structures called gemmae, which are bundles of cells that reside in cups on the surface of the plant and are dispersed by raindrops.  Their rhizoids are composed of single, elongated cells, not multiple cells as in moss. Anthocerophyta The third and final division, Anthocerophyta, consists of approximately one hundred species of hornworts.  These plants resemble liverworts in their gametophytes, but they can be easily distinguished by their sporophytes.  Hornworts possess elongated capsules that grow like horns from the gametophytes, arising from a group of cells at the base of the horn that divide continuously throughout the sporophyte's life span.  This feature is unique among plants: hornworts are the only known plants to posses a continuously dividing group of cells.  Another unique feature of hornworts is their photosynthetic cells: each possess a single large chloroplast instead of the many smaller chloroplasts that most plants have.

Societal Impact

Due to their limited range of habitats, nonvascular plants have not had that great of an impact on society.  Because of their sensitivity to the world around them, though, they can be useful indicators of environmental conditions. For example, some mosses are strict calcicoles, and they will only grow where calcium is freely available in the substrate.  Nonvascular plants are also particularly susceptible to air and water pollution, which makes them good indicators of the purity of the environment.  For example, in the countryside, trees generally have large numbers of mosses and liverworts growing on them, but in towns and cities the trees are generally bare.  Mosses are also used in a lot of decorative gardening, and they also serve as food for small animals and insects.


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