Alternation of Generation between Mosses and Ferns
Literature Review Chapter Introduction
This section will be used to introduce the chapter which presents a review of the relevant scholarly and peer-reviewed literature to provide a general background concerning the alternation of generation between mosses and ferns and the specific life cycles for these organisms. A brief summary of the research will conclude this chapter.
Background Information about Mosses
Mosses are part of the phylum, Bryophyta. Mosses do not have vascular tissue, are invariably small and tend to grow in wet or very moist areas; there are about 14,000 known species mosses (Hanson 1991:28). Some moss species have commercial value, such as sphagnum peat moss, which is described by Arcand and Talbot as “a natural organic material created in very humid conditions when plant debris. Peat moss in its raw state is a spongy, fibrous, lightweight material whose color varies from light to dark brown depending on the degree of decomposition” (2000:36). Both mosses and ferns are dependent on the presence of lichens to help establish a foothold in rocky outcroppings where lichens form soil deposits (Oplinger, Halma and Halma 2006). According to Oplinger and his associates, “Only then can mosses and ferns assume a role in the succession process” (2006:24). Based on their analysis of the evolution of bryophyte sequences, Floyd, Zalawski and Bowman report that, “The ancestors of moss can be inferred to have inherited a single gene from an algal ancestor [and] the conducting tissues of mosses may be either analogous or homologous to those of vascular plants” (2006:373).
Background Information about Ferns
Ferns are commonly found in forest areas; these organisms have sporophytes that are typically delicate in appearance with broad leaves that are highly divided (Mohlenbrock 2003). Ferns that prefer moist, moss-covered soil on rock outcroppings include berry bladder fern and brittle bladder fern (also called fragile fern); both of these species form spores as well as creating asexual “bladders” which are pieces of tissue that are capable of giving rise to a new plant (Mohlenbrock 2003). By contrast, walking fern do not resemble many other types of ferns with leaves that are undivided and narrow (Mohlenbrock 2003). This species is capable of reproducing in such a fashion that it can “walk” across rocky outcropped surfaces that contain sufficient amounts of suitable soil (Mohlenbrock 2003). Other species of fern, including purple cliff-brake, two species of Woodsia, and the spike moss northern selaginella, prefer drier environs (Mohlenbrock 2003). Interestingly, farmers in China used burned ferns (primarily Dicranopteris dichotoma) as a crop fertilizer (Coggins 2002) and ferns are the most diverse component of the flora of the Southern Pole region (Falcon-Long, Cantrill and Nichols 2001).
Alternation of Generation in Mosses
According to McDaniel, Willis and Shaw, “Like all land plants, the moss life cycle consists of a multicellular haploid gametophyte generation that alternates with a morphologically distinct diploid sporophyte generation” (2007:2489). Likewise, Polunin reports that, “The mosses are all rather small plants in which the gametophyte, during the greater part (but by no means all) of its life, consists of a more or less upright stem bearing small leaves” (1960:51). The various stages of the life cycle of moss are described by McDaniel and his colleagues thusly: “The gametophyte generation consists of pollen and ovules, while the sporophyte generation is much larger and long lived. In contrast, the gametophyte is the dominant portion of the moss life cycle, and the sporophyte is ephemeral” (2007:2490). In addition, McDaniel, Willis and Shaw add that, “Recombinant haploid spores are produced in a single cross between two moss gametophytes” (2008:1425).
Alternation of Generation in Ferns
Ferns are vascular plants that do not develop seeds as a function of their reproductive cycle (Mohlenbrock 2003). According to Mohlenbrock, “Like many plants, their generations alternate between a spore-producing form, called the sporophyte, and a gamete-producing form, called the gametophyte. In vascular plants, the sporophyte is the plant people usually see and recognize” (2003:56). In ferns, sporophytes result in haploid cells containing just one from each pair of parental plant chromosomes, and these cells are known as spores (Mohlenbrock 2003).
Following dispersal, the spores give rise to gametophytes which then produce sex cells known as gametes (Mohlenbrock 2003). New sporophytes can be produced when two gametes combine and restore the double number of chromosomes (Mohlenbrock 2003).
This section will be used to provide a summary of the research and important findings supported by citations.
Arcand, Yves and Pierre Talbot. (2000) “Using Peat to Treat Wastewater.” Journal of Environmental Health 62(6): 36.
Author provides a description concerning the utility of peat moss in wastewater treatment applications. Included in the report is a discussion concerning the origins of habitats favored by peat moss. Author also presents a useful basic description of this species and its physiology.
Coggins, Reed. (2002). “Ferns and Fire: Village Subsistence, Landscape Change, and Nature
Conservation in China’s Southeast Uplands.” Journal of Cultural Geography 19(2): 129-
Author describes the economic impact of fern species for agricultural practices in rural
China where ferns are burned and used as fertilizer as well as used as a rotation crop to improve crop yields. Report provides useful background information for ferns.
Falcon-Long, H.J., D.J. Cantrill and C.J. Nichols. Biodiversity and Terrestrial Ecology of a Mid-cretaceous, High-latitude Floodplain Alexander Island, Antarctica. Journal of the Geological Society 158: 709-711.
Authors present their findings concerning the flora of Alexander Island, Antarctica,
including several species of ferns and the role they play in the region’s biodiversity.
Authors include several descriptions of various life stages of ferns as well that will provide useful information for the proposed study.
Floyd, Sandra K., Christopher S. Zalewski and John L. Bowman. (2006). “Evolution of Class Iii
Homeodomain-leucine Zipper Genes in Streptophytes.” Genetics 173(1): 373-374.
Authors examine evolution of various land-based plants to determine the developmental roles of a specific gene family and found that as land plants became more complex through gene duplications, so too did the role of the investigated gene family, class III
HD-Zip. Authors identify future study directions for ferns that may provide a useful point of departure for further analysis.
Hanson, Earl D. Understanding Evolution. New York: Oxford University Press, 1991.
Author presents a comprehensive overview of mosses and ferns as well as their reproductive cycles. Although the focus of this text is on the role played by evolution in promoting modern species, author provides a valuable straightforward description of these organisms and how they function in the environment.
Mohlenbrock, Robert H. (2003, October). Fern relations. Natural History, 112(8): 56-57.
In this study, author describes the flora of southwestern Massachusetts including several species of ferns, as well as their reproductive and life cycles that are useful additions to this literature review. Vivid descriptions of various fern species are supplemented by the growing environment preferred by different species. Study provides useful background information and specific data concerning life cycles of ferns that will be valuable additions to the proposed study.
Oplinger, Carl S., Robert Halma and Robert Halma. The Poconos. New Brunswick, NJ: Rutgers
University Press, 2006.
Authors review the flora and fauna of the Poconos, including a comprehensive discussion of the life cycles of various species of ferns and mosses. While their discussion and analysis is restricted to the Poconos region, there is some useful guidance concerning the alternation of generation between mosses and ferns that will be useful for this study.
McDaniel, Stuart F., John H. Willis and a. Jonathan Shaw. (2008). “The Genetic Basis of Developmental Abnormalities in Interpopulation Hybrids of the Moss Ceratodon
Purpureus.” Genetics 179(3): 1425-1426.
In this follow-up study to the 2007 study listed below, authors present the results of their research into the genetic basis of abnormal hybrid development in the offspring of crosses between two isolates of the moss, Ceratodon purpureus. As part of their background information, authors present a useful description of mosses and their life cycles that will be a valuable addition to the outlined study.
McDaniel, Stuart F., John H. Willis and a. Jonathan Shaw. (2007). “A Linkage Map Reveals a Complex Basis for Segregation Distortion in an Interpopulation Cross in the Moss
Ceratodon Purpureus.” Genetics 176(4): 2489-2491.
The purpose of this study was to identify segregation patterns across the genome in general and sex chromosomes in particular of a species of moss. As part of their background information, authors present a useful description of mosses and their life cycles that will be a valuable addition to the outlined study.
Polunin, Nicholas. Introduction to Plant Geography and Some Related Sciences. London:
This biology text has several sections devoted to Byrophtya, including the mosses, including detailed descriptions of their life cycles and the processes that are involved.
Author also provides several graphics that will be useful in illustrating these life cycle processes for mosses.
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