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The Eukaryote life-cycle—diploidy, haploidy & sex Eukaryotes •Have membrane bound organelles eg chloroplasts, mitochondria of prokaryote origin (endosymbiosis) •Larger and more complex genetic (multiple chromosomes) and membrane systems than prokaryotes •Usually with some diploid stages, and often sex somewhere in their life-cycle The Eukaryote life-cycle—diploidy, haploidy & sex Although sex is often associated with diploidy, and fertilization (syngamy) produces a diploid cell, only haploid cells can combine sexually (become gametes) meiosis is required to restore the haploid state meiosis restores the haploid state but also generates infinite genetic variety among the pool of gametes that it forms Where is the life-cycle does meiosis take place? What advantages/disadvantages does diploidy bring? What are the advantages/disadvantages of sex? Meiosis reduces chromosome number from diploid to haploid: a closer look • Many steps of meiosis resemble steps in mitosis. • Both are preceded by the replication of chromosomes. • In meiosis, diploid cells divide twice in a row after replicating their chromosomes only once. • Each final daughter cell is haploid—one copy of each chromosome • In mitosis cells divide but ploidy doesn’t change • Haploid cells divide to form haploid cells and diploid cells divide to form diploid cells • Meiosis reduces chromosome number by copying the chromosomes once, but dividing twice. • The first division, meiosis I, separates homologous chromosomes. • The second, meiosis II, separates sister chromatids. Fig. 13.6 Sexual life cycles produce genetic variation among offspring • Three mechanisms contribute to genetic variation: – independent assortment of chromosomes (2n combinations where n=haploid number) – random fertilization—any sperm can fertilize any ovum – crossing over—each gamete is unique • By independent assortment alone each individual chromosome in a gamete would be exclusively maternal or paternal in origin. • However, crossing over produces recombinant chromosomes which combine genes inherited from each parent. Fig. 13.10 Diploidy vs Haploidy—eukaryotes usually have both in their life cycle Diploid cells have two copies of their entire genome •provides genetic redundancy which increases resistance to mutagens like ultraviolet light •having two copies allows species to benefit from sex(genetic exchange) with other individuals—two copies different. This requires meiosis •however cell cycle and cell division more complex Haploid cells have only one copy of their genome •more vulnerable to genetic damage •cell cycle & cell division less complex •can sometimes function as gametes—combine with other haploid cells–syangamy or fertilization Meiosis restores the haploid condition (2n to n) can take place at different stages of the life cycle in the zygote (zygotic meiosis) later, after a diploid organism (sporophyte) has developed sporic meiosis (gametophyte (n) & sporophyte (2n)) gametic meiosis (no gametophyte) Zygotic meiosis is the simplest sexual life cycle displayed by eukaryotes. Why? Eg Chlamydomonas—a unicellular green alga Chlamydomonas: the sexual life-cycle resistant zygospore Gametes are of two mating types, + and ─ We call this type of life-cycle zygotic meiosis, why? This is the simplest type of sexual life-cycle found in eukaryotes? Explain? http://academic.kellogg.cc.mi.us/herbrandsonc/bio111/algae.htm The life-cycle of Oedogonium: a different version is found in filamentous algae. In what way is this life-cycle similar to that of Chlamydomonas, and how is it different? Because Oedogonium is a multicellular organism (filamentous) its life cycle allocates asexual growth potential to both increase in filament length (mitotic divisions within filaments) and to increase in filament number (zoospore formation). In contrast, since Chlamydomonas is a unicellular organism, all mitotic divisions result in more new organisms http://academic.kellogg.cc.mi.us/herbrandsonc/bio111/algae.htm Oedogonium: unbranched, filamentous cylindrical cells,distinctive rings at the apical ends of certain cells. Each cellular division creates a new ring on the cap cell. Cells contain a parietal, netlike chloroplast with several pyrenoids. asexual reproduction by zoospore; sexual reproduction by fertilization by egg and sperm; Germination of a zoospore Vegetative cells Apical rings http://protist.i.hosei.ac.jp/pdb/images/Chlorophyta/Oedogonium/sp_10c.html Oedogonium: a mat-forming filamentous green alga http://images.google.ca/imgres?imgurl=http://www.csupomona.edu/~jcclark/classes/bot125/resource/graphics/g/chl_oedogonium.jpg&imgrefurl=http://www.csup omona.edu/~jcclark/classes/bot125/resource/graphics/chl_oedogonium.html&h=298&w=450&sz=20&hl=en&start=11&tbnid=oTj8oIOpG56qM:&tbnh=84&tbnw=127&prev=/images%3Fq%3Doedogonium%2B%26svnum%3D10%26hl%3Den%26lr%3D%26sa%3DN Oogonium antheridium Class Charophyceae Chara A green macrophytic alga with a slighly different zygotic type of life cycle. Meiosis occurs within the zygospore, which germinate into haploid plants that produce oogonia and antheridia What is the main difference between the Chara and Oedogonium life-cycles? Charophyceans are considered to be closely related to land plants Chara sperm An oogonium with an open antheridium http://images.google.ca/imgres?imgurl=http://www.visualsunlimited.com/images/watermarked/316/316013.jpg&imgrefurl=http://ww w.visualsunlimited.com/browse/vu316/vu316013.html&h=238&w=350&sz=12&hl=en&start=13&tbnid=Suqmn7RgKRdyIM:&tbnh=78&t bnw=116&prev=/images%3Fq%3Dchara%2Boogonium%26svnum%3D10%26hl%3Den%26lr%3D%26sa%3DG Spirogyra A filamentous green alga with a long spiral chloroplast A peculiar form of sexual reproduction— conjugation to form zygospores, with no motile gametes or zoospores Meiosis again takes place in the zygote stage: haploid cells grow into new filaments (asexualy)—no motile zoospores Filaments are either male or female mating types During sex, when male and female filaments are lined up next to each other, the male type cytoplasm migrates through the conjugation tube and joins with the female cytoplasm to form a zygote—no motile gametes. Spirogyra filaments produce a thin layer of slime that tends to keep them clean of epiphytes, why would this be of particular importance to them? http://academic.kellogg.cc.mi.us/herbrandsonc/bio111/algae.htm Ulva—sporic (biphasic)—isomorphic alternation of generations What is the most fundamental difference between zygotic and sporic meiosis cycles? What advantage might sporic life cycles have overy zygotic cycles? Steps in the Cladophora Life Cycle •Quadriflagellate zoospores (1N) form in ordinary vegetative cells of the sporophyte individual (2N), break through the sides of the cells and enter the water •Spores settle on substrate and germinate into diecious gametophytes (1N) •Gametophytes form biflagellate isogamous gametes (1N), which burst from lateral apertures in branches of gametangia •Gametes fuse externally forming quadriflagellate zygote (2N) •Zygote settles and germinates directly into Sporophyte (2N) Cladophora: filamentous benthic green alga with branched filaments 500 microns Advantages and disadvantages of a biphasic life cycle If the biphasic cycle is heteromorphic such as in kelp or red algae, then the gametophyte, and sporophyte might be adapted to different ecological niches. This can be advantageous if both types of niches occur commonly enough to allow the life cycle to be completed. Eg. the two phases often differ in terms of size, vulnerability to waves action, and to predators. If they do not, and say only the sporophyte or the gametophyte is successful, then the sexual life-cycle could not be completed. There are many asexual algae which may have lost their sexuality for this reason. There are some examples of brown algae where the gametophyte is lost, and meiosis in the sporophyte gives rise directly to gametes rather than spores (gametic meiosis). These species are usually intertidal—that is exposed to light and drying during the tidal cycle, to which the gametophyte may be too sensitive. If the biphasic cycle is isomorphic, the cycle would make little sense unless there were subtle differences between the sporophyte and the gametophyte. Several studies have found differences in growth rate, nutrient requirements, vulnerability to stresses and ability to proliferate asexually. Zygotic meiosis—the simplest sexual life cycle, common among fungi and protists. – The zygote is the only diploid phase. – After fertilization, the zygote undergoes meiosis to produce haploid cells. – These haploid cells undergo mitosis to either: • increase in number • or grow into a haploid multicellular adult organism. – Some haploid cells develop into gametes by mitosis. Sporic meiosis—alternation between haploid and a diploid organisms—diploid zygote first replicates by a series of mitotic divisions to form a diploid organism, which undergoes meiosis. (some green and brown algae, red algae and plants) – both haploid (gametophyte) and diploid (sporophyte) multicellular stages. – Meiosis by the sporophyte produces haploid spores that develop by mitosis into the gametophyte. – Gametophyte produces gametes by mitosis. Fertilization forms a zygote which produces a sporophyte by mitosis The timing of meiosis and fertilization varies among species. Gametic meiosis—gametes the only haploid stage found in all animals incl. humans & some protists. – haploid cells don’t divide, – diploid zygote that divides by mitosis to produce a multicellular organism. Fig. 13.5a Question What is the difference between a spore and a gamete? Both are reproductive cells, but •a gamete must combine with another gamete of the opposite sex •a spore can produce a new multicellular organism by mitosis all by itself (no sex required) http://academic.kellogg.cc.mi.us/herbrandsonc/bio111/algae.htm