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H 8 0޽h ? ̙330 pDB(  D^ DS     Dc $_ @   8 9hncNucTb[P[ṽNegnbNTR N TSR:N T[MT̃SOꁫSU[ N+RṽSO.^R rzۏL g'`uk0 S_ TÑSOv$N9h̃Nu TÑNvRgvNce OSNucTb[P[0 _[MT̃SOꁫ NU[ PR+RvSNT̃SOv N TNMWegۏL g'`uk0sS[$Ny N T̃|ṽNvGMbb_bcTb[P[0 ُ$Ny N T̃|ṽSO(Wb_`0'Y\ NN,e:S+R FOut N g]+R 8^(u + T - egh:y0YgNỹ|bMP[V:N + HN0Q/fNKNcT b_bcTb[P[vSÑ|bMP[V:N - 0 5 cTb[P[v S QpeRˆTb_b4*NUS PSOv8h cTb[P[ Se N*N8hۏeQN*N{ {Sbb[P[V 8h~Y!k gNRˆ(Wb[P[VQNu@b gv8h02bkяNA~k0 Sexual reproduction in the Zygomycetes: There are two possible nuclear states in the mycelia of this group of fungi. They can have a single type of nucleus in their mycelium, a condition termed termed homothallism, or they can contain the two mating type nuclei within their mycelium, termed heterothallism. If the fungus is homothallic the first event in the onset of sexual reproduction has to be somatic fusion. This is termed conjugation. To achieve such a mating it is necessary to attract each other and an elaborate sequence of cellular and biochemical events have been established for some of these fungi. This signalling involves the secretion of inducer molecules that are responsible for causing the formation of zygophores, modified hyphal tips, and these then grow towards each other long a gradient of hormone. The exact sequence is shown below. Sexual reproduction in the Zygomycetes: Once in contact the two zygophores fuse, and then the nuclei fuse to form the diploid. Meiosis occurs, producing four haploid nuclei, but three may degenerate. The timing of fusion varies from species to species. &Zygomycete Asexual Reproduction Zygomycete fungal mycelium is coenocytic. At the onset of sporulation large amounts of aerial hyphae are produced. The tips of these aerial hyphae fill with cytoplasmic contents, and the nuclei undergo repeated mitosis. Around each of the nuclei cytoplasm and organelles collect, and by the formation of copious vesicles from the Golgi, each nucleus becomes isolated from the next by a plasma membrane. Within the spaces created by this cytoplasmic cleavage, spore walls begin to form, again by the fusion of Golgi vesicles containing cell wall monomers and enzymes with the spore membrane. A sporangium forms. As these events occur so there is considerable water uptake by the forming sporangium, and as the columella forms the structure comes under considerable turgor pressure. The large sporangia can contain up to 100,000 spores (Fig 3). 2 [ 2  2!%fA!%f!R$'5   CG  p  =        (  f = .KH D 0޽h ? ̙33L 0  L(  L^ LS     Lc $L @   9hncNucTb[P[ṽNegnbNTR N TSR:N T[MT̃SOꁫSU[ N+RṽSO.^R rzۏL g'`uk0 S_ TÑSOv$N9h̃Nu TÑNvRgvNce OSNucTb[P[0 _[MT̃SOꁫ NU[ PR+RvSNT̃SOv N TNMWegۏL g'`uk0sS[$Ny N T̃|ṽNvGMbb_bcTb[P[0 ُ$Ny N T̃|ṽSO(Wb_`0'Y\ NN,e:S+R FOut N g]+R 8^(u + T - egh:y0YgNỹ|bMP[V:N + HN0Q/fNKNcT b_bcTb[P[vSÑ|bMP[V:N - 0 z [ 2#'fA#'f# H L 0޽h ? ̙33 0 NFT(  T^ TS    @ Tc $ r @   d0P[Vb[P[ (WP[VQb_bv g'`b[P[0b_bP[Vb[P[/fP[Ṽ~v;Nyr_0 P[V$N'`~ހcNTb_bvVr~g0P[V gtb_0hb_0WR{b_0eb_I{ Vy _0 P[VQb[P[8^/f1 8*N0 P[Vb[P[vb_r0'Y\0r_NT Nv T0 N TvP[Ṽb_bP[Vve_ N T g{USv/f$N*N%{Q~ހ~Tb_bP[V ~ހ8hRˆb_bP[8h kNP[8hb_bN*NP[Vb[P[0ORuk0B 1 3'f# H T 0޽h ? ̙33 0 \`(  \^ \S     \c $($ @   VtP[Ṽb_bP[V'Ye_T_ g{USv/f$N*N%{Q~ހcTTvcb_b YRukؚI{w̃b_bP[Vv$N'`~ހ] gNRS b_` N_N gf>fv:S+R0  ̖'`vyNVhVascogonium);Ė'`vyĖhV(antherdium)0   (WP[VTP[Vb[P[SǏ z-N,ĖhVNNVhV Nbv~ހuQỸN,v^ gĉ_0W\NVNSVOO,b_bP[Vgascocarp)0P[Vg;N g Ny{|W0 ̃ N T'`+RṽN RSQĖhV\ TNVhV'Y $N*N'`hV[cT ĖhVvQ+TirǏS|NۏeQNVhV ۏL(M0(MT NVhVuQYw̃NyNVN NVNvzv~ހ/fS8hv (Wvz~ހQSu8hM bP[Vk~ހ0Q~ gNRˆTQpeRˆNu1-8*NP[Vb[P[0 P[Vg(WP[VTP[Vb[P[SǏ z-N ĖhVT̖hV Nbv~ހuQỸN b_bOb~~ te*N~gb:NNP[[SO0ُy g'`vP[[SOy:NP[Vg P[VS(WvQ-N0 P[Vg;N g Ny{|WNy:N[hQ\_ yVX0tb_ gT[SvyP[VX0_SHTvrvyP[Vv0 B #|N  3!o |H \ 0޽h ? ̙33 0 WOd(  d^ dS    I dc $8 @   Figures F, G. Ascomycota. Sexual reproduction in this group leads to the production of one or more asci, each containing usually 8 ascospores. Figure G shows a cluster of asci of Sordaria macrospora; the mature ascospores are black. Often the asci are contained in a fruiting body which may be flask-shaped (termed a perithecium and shown for S. macrospora in Figure F) or cup-shaped (an apothecium) or closed (a cleistothecium). See Lichens for examples of apothecia; see Thermophilic microorganisms for an example of a cleistothecium " K   F      !0!0H d 0޽h ? ̙33 0 l$(  l^ lS     lc $P @    Sexual reproduction in the Ascomycetes: In this group of fungi there are no specialized organs of hyphal fusion, different mating type mycelia merely fuse with each other to form transient dikaryons, mycelia with two mating type nuclei within it. The dikaryotic mycelium can differentiate to from varying amounts of sterile mycelium around what is to become the fertile tissue of the fruit body. In yeasts, a single, diploid yeast will undergo meiosis, producing four haploid progeny cells, but in more complex fungi there are a sequence of cellular and nucleic events that ensure an organized fertile layer. The events are illustrated below in figure Fig. 8. &Spores are delineated around these nuclei in a process called free cell formation, and as most of the cytoplasm is contained around the nucleus and within the spore wall, all that is left outside is cell sap. These modified hyphae are termed Asci, and the spores that are held within them are termed ascospores. The asci are often found packed tightly with other asci, and between a dense layer of supporting sterile tissue. Often the structure is large enough to be seen with the naked eye. &The asci can be aggregated together in various sorts of fruit body which we will see in the practical, including the, cup fungi (Discomycetes, apothecial), the flask fungi, (Pyrenomycetes, perithecial), the mildews (Plectomycetes cleistothecial) and the fungi with black, crusty stromata (Loculoascomycetes, pseudothecial fungi). There are also the yeasts, Hemiascomycetes,. Their ascospores are normally formed in loose asci and are not actively discharged. We have not looked at these. When they form ascospores in fruit pulps or liquids they are usually liberated by the disintegration of the ascus wall. d' K <3 <U 5 p 6 +y !  # $  N SH l 0޽h ? ̙33$0 xt(  x^ xS     xc $t+ @l  TL___PPT9.& Sexual reproduction in the Basidiomycetes: Basidiomycetes are characterised by the most complex and large structures found in the fungi. They are very rarely produce asexual spores. Much of their life cycle is spent as vegetative mycelium, exploiting complex substrates. A preliminary requisite for the onset of sexual reproduction is the acquisition of two mating types of nuclei by the fusion of compatible mycelium. This creates a dikaryon where single copies of the two mating type nuclei are held within every hyphal compartment for extended periods of time. Maintenance of the dikaryon requires elaborate septum formation (clamp connections, Fig 10.) during growth and nuclear division. O&nset of sexual spore formation is triggered by environmental conditions and in the larger Basidiomycetes begins with the formation of a fruit body primordium. The primordium expands and differentiates to form the large fruit bodies of mushrooms and toadstools. The mycelium within this structure remains as a dikaryon, diploid formation only occurring within the modified hyphal tip called the basidium. Meiosis occurs within the basidium, and the four products are extruded from the tip of the basidium on sterigma (below). Usually this event occurs across a large area of basidia called a hymenium, or fertile layer. It is usually formed over an extensive sterile layer of tissue like a mushroom gill. &There are three major divisions in the basidiomycetes. Hymenomycetes. Basidia are in extensive fertile layer which are susceptible to rain when exposed. Spores are actively discharged from a protected hymenium when ripe. This group includes mushrooms and toadstools, boletes, brackets and coral fungi. Gasteromycetes. Hymenia line closed cavities in an initially closed fruit body. Basidiospores are released passively by autolysis of the hymenium, and basidiocarps disintegrate at maturity. This group includes earth balls, puff-balls, stinkhorns and birds nest fungi. Teliomycetes. These are the rusts and smuts, neither of which form large, conspicuous, fruit bodies but invade plants and produce characteristic sporulating lesions in plant tissue. 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