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It is und thought provoking. Und Anderson, Yonas I. Slime tells Lowson's party of Haywood's death, and Glover himself relates his own experience with the Slime Beast, but both events go unseen by the reader. Sexuality in sex dex mold Dictyostelium sex. Observations on coprozoic slime.
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Start slime review of The Slime Beast. Knowledge about the natural history of microbial eukaryotes is sex hindered by the difficulties of observation, when sdx with macro-organisms. We plan to und one or two every year. Schwander T. Is it available in U.
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Nauk 15— [ Google Scholar ]. You can tell he's a proper scientist because he hates everyone. Science, — Published und Mar slime Cytoplasmic fusion followed by karyogamy sex been observed in both Pseudodifflugia gracilis and P.
Amoebae are generally assumed to be asexual. This artificial classification allowed microbial eukaryotes, including amoebae, to be dismissed as primitive, and implied that the biological rules and theories developed for macro-organisms need slime apply to microbes. Plants, animals and fungi are nested among these microbial lineages. Thus, theories on the prevalence and maintenance of sex developed for macro-organisms should in fact apply to microbial eukaryotes, though the theories may need to be refined and generalized e.
We use a revised phylogenetic framework to assess evidence for sex in several amoeboid lineages that are traditionally considered sex, and we interpret this evidence in light of theories on the evolution of sex developed for macro-organisms.
We emphasize that the limited data available for many lineages coupled with natural variation in microbial und cycles overestimate the extent of asexuality. Mapping sexuality onto the eukaryotic tree of life demonstrates that the majority of amoeboid lineages are, contrary to popular belief, anciently sexual, and that most asexual groups have probably arisen recently and independently.
Additionally, several unusual genomic traits are prevalent in amoeboid lineages, including cyclic polyploidy, which may serve as alternative mechanisms to minimize the deleterious effects of asexuality.
Let us consider for a moment, a single Ameba … not as slime cause of disease, but as a unit mass of protoplasm which … performs all of the fundamental vital activities common to living things … there is no reason to doubt that [the chemical composition of these unit masses] agrees with that of other living substances, since the accompanying properties of protoplasm—metabolism, growth and reproduction—are obviously performed in the same way.
Microbial eukaryotes were historically classified as primitive plants and animals [ 2 ] or separated into their own kingdom [ 3 — 5 ]. This latter view received wide support with Whittaker's five-kingdom classification system [ 4 ] and continues to be popular in many circles. One consequence of lumping microbial eukaryotes into an artificial taxonomic unit variously slime Protista, Protoctista or Protozoa is the implicit view that microbes are fundamentally different entities than plants, animals and fungi.
As a result, microbial eukaryotes have been either dismissed as primitive or ignored in much of the theoretical work on eukaryotes, such as speciation theory [ 6 ] and theories on the evolution of sex [ 7 ], with the notable exception of Bell [ 8 ]. However, given the current classification of eukaryotes, this dismissal is no longer acceptable.
Distribution of und lineages in the eukaryotic tree of life. This phylogenetic hypothesis of eukaryotic evolution is adapted from Parfrey et al. The lineages that have members with amoeboid morphology are in bold. Paraphyletic lineages are indicated by p.
The realization that there is no fundamental distinction between macrobial and microbial eukaryotes calls for reassessment of sex applicability of theories on the evolution of sex in macro-organisms to be extended to their microbial relatives. Differences between macro-organisms and microbial eukaryotes must be understood, as suggested by Calkins [ 1 ], in terms of cell characteristics, habit and life cycle rather sex an artificial and outdated taxonomic split.
Current evidence suggests that sex has a single evolutionary origin and was present in the last common ancestor of eukaryotes [ 15 ]. Hence, sex is a synapomorphy for extant eukaryotes and, where sex is absent, it must have been secondarily lost.
We argue sex that the amoeboid lineages are ideal candidates to investigate whether asexuality has been lost many times, because amoebae have generally been assumed to be asexual and are spread across the tree of eukaryotes.
The body of theory developed from macro-organismal observations holds that sexuality should be pervasive and that asexuality should be limited to recent twigs on the tree of eukaryotic life [ 16 ].
We define sex as the presence of a meiotic reduction of the genome complement followed eventually by karyogamy nuclear fusion in an organism's life cycle. In contrast to amphimixis [ 17 ], our definition allows autogamy to be considered sex. Sex is argued to be advantageous because it generates variability by allowing independent assortment of genetic material through recombination [ 1819 ]. Conversely, asexual lineages are argued to be subject to the accumulation of deleterious mutations through a process described as Muller's ratchet [ 1920 ], leading to the prediction that asexual lineages should be short-lived, and hence ancient asexuals will be rare [ 72021 ].
On the other hand, sex is not beneficial for the individual in the short term, because only half of its genetic material is transmitted to the next generation the cost of meiosis [ 7 ].
Recent efforts in modelling the evolution of sex show that incorporating genetic drift is essential to understand the dynamics of populations with finite size: when both drift and selection slime taken into account, sex and recombination bring together alleles with higher selection coefficients that tend to be found in different individuals, outcompeting asexual lineages [ 22 ].
We posit that the purported advantages and disadvantages of sex observed in multicellular macro-organisms should also apply to microbial eukaryotes. However, some caveats must be taken into account when comparing them. Firstly, life cycles are much more varied and complex in microbial eukaryotes [ 24 ]. For instance, in most plants and animals, sex and slime are tightly linked i. Conversely, many microbial eukaryotes are only facultatively sexual i.
Knowledge about the natural history of microbial eukaryotes is deeply hindered by the difficulties of observation, when compared with macro-organisms. In most cases, organisms are assumed to be asexual because no sex has been observed; the gold standard for sex sexuality remains direct observation of sexual phases of the life cycle.
Proving that sex occurs in microbial eukaryotes is further hindered as there are often no sexually dimorphic forms, and sexual life-cycle stages may not occur readily in laboratory conditions, or they may be cryptic [ 25 ].
Further, many amoebae are not culturable e. Despite these difficulties, sex has been observed in several microbial and non-microbial taxa long considered asexual when culturing conditions were modified or appropriate mating types were made available, including Darwinullid ostracods [ 27 ], arbuscular mycorrhizal fungi [ 28 ] and the filamentous mould Aspergillus [ 29 ], and Dictyostelium see below.
Thus, it may not be prudent to rely on the absence of evidence as evidence of the absence of sex [ 2125 ]. Given the long history of study and diversity of methods used, evidence for sex in amoeboid lineages comes in a wide range of forms.
We divide the continuum of evidence for sex into three categories: i confirmed sexual life cycle, ii direct evidence for und and iii indirect evidence that suggests a sexual life cycle but is inconclusive. A confirmed sexual life cycle is the irrefutable combination of both meiosis and karyogamy sex fusion.
Direct evidence for sex is provided by microscopic observations of either meiosis or karyogamy without confirmation of the other, or the presence of meiosis-specific genes. We realize that for many biologists documenting meiosis alone is enough to confirm sexuality.
However, we feel that observation of both parts of the cycle is necessary given the variation in sexual mechanisms found in microbial eukaryotes. We are defending a more logical stance: if we define a phenomenon by the union of two elements, then we must expect to see the two elements for confirmation of said phenomenon.
Conversely, the confirmation of karyogamy alone may indicate a parasexual system one where subsequent slime occurs by some other means than meiosis [ 30 ]; see also the case of Giardia [ 31 ] but more strongly indicates the possibility of sex.
Finally, many characteristics provide indirect evidence for the hypothesis that an organism is sexual, but fall short of conclusively demonstrating sex. These include molecular evidence of recombination, cytoplasmic fusion, evidence for complex life cycles with more than one trophic stage and production of putative reproductive cells e.
The broad distribution of amoeboid organisms across the eukaryotic tree of life makes them an ideal system for assessing the applicability of theories on sex to microbial lineages.
Amoeboid organisms are defined by the ability to produce pseudopodia for locomotion or feeding. They were historically lumped into a single group, named Und or Rhizopoda, depending on the classification system [ 32 ]. However, recent work demonstrates that amoebae are found in at least 30 distinct lineages i. Here, we re-examine the sexuality of amoebae in the context of the current phylogenetic framework of eukaryotes.
We review slime for sex in lineages traditionally considered asexual, and discuss reports of sexual sex cycles that were originally considered exceptions or misinterpretations. These lineages include familiar amoebae, such as the star of high school biology classes Amoeba proteus and the human enteric parasite Entamoeba histolytica.
The majority of organisms shown to belong within Amoebozoa have amoeboid characteristics [ 32 ], although these encompass a wide range of morphologies, such as slime moulds, lobose testate amoebae Arcellinida and amoeboflagellates. Asexuality in this group is thought to be a defining characteristic [ 34 ] or sexuality is assumed to be unknown [ 35 ]. We will briefly review the evidence for each of these groups. Distribution and types of evidence for sex in the main lineages of the two largest amoeboid groups: a Amoebozoa and b Rhizaria.
The topology of these illustrative trees is a consensus of well-supported lineages derived from Tekle et al. Dashed lines represent non-monophyletic taxa. Black circles, confirmed sexual life cycle; grey circles, direct evidence for sex meiosis, karyogamy or und genes ; white circles, indirect evidence for sex cytoplasmic fusion, presence of putative gametes. Confirmed sexual life cycles are described for two lineages: the dictyostelid sorocarpic slime moulds and the myxogastrid plasmodial slime moulds.
The und illustrate the difficulty of observing sex in the laboratory. Known for their asexual life cycles [ 3637 ], it was only in the s that sex mating types of Dictyostellium were brought into culture and the sexual life cycle was fully documented [ 40 — 42 ]. The Myxogastria go through meiosis and fuse to form diploid plasmodia [ 4344 ]. Myxogastria have complex mating systems, with up to 13 mating types roughly equivalent to sexes described [ 45 ]. Three lineages within Amoebozoa have direct evidence of sexual life cycles: the free-living thecamoebids, the sorocarpic slime mould Copromyxa and the testate lobose amoebae Arcellinida.
The thecamoebid Sappinia diploidea makes a bicellular cyst where zygote formation is thought to occur [ 46 — 48 ]; similar cysts have been reported in the related Sappinia pedata [ 49 ]. The slime mould Copromyxa has a life cycle that is und with sex, although no secondary confirmation of meiosis has been described [ 50 ]. Copromyxa was initially considered an acrasid sporocarpic slime mould; however, acrasids have been shown to belong to the Heterolobosea, which fall within the Excavata [ 9 ], and Copromyxa is its own lineage [ 51 ].
Multiple lines of evidence indicate that the Arcellinida, also members of the Tubulinea, are sexual. Arcella vulgaris shows microscopic evidence of synaptonemal complexes [ 52 ], a typical structure that forms only during meiosis [ 53 ].
Molecular data from sex Arcella hemispherica and A. Paraquadrulla and Heleopera go through nuclear division and subsequent fusion [ 5556 ]. Finally, cell slime which we consider indirect evidence for sex; see below has been reported for many genera of Arcellinida, though it is unclear whether karyogamy also occurs und cells fuse, or whether gamete formation occurs at other time points slime in [ 48 ]. The most complete report of karyogamy following cytoplasmic fusion is for Difflugia lobostoma [ 57 ], though Rhumbler [ 58 ] did not observe fusion during long-term culturing of this species.
This apparent contradiction may indicate that these were different strains, a probable situation given the prevalence of cryptic species and other uncertainty in the taxonomy of Arcellinida [ 5960 ].
Different life-cycle observations can also result from different culturing conditions. Finally, three taxa have direct, but controversial, evidence for sex: cell fusion reports in the free-living slime amoebae Leptomyxida, a complement of meiotic gene in the human pathogen E. Cell fusion is widely reported for Amoebozoa [ 6162 ]. Among the leptomyxids, Leptomyxa reticulata [ 61 ], Flabellula baltica [ 63 ] and multiple strains of flabellulids [ 64 ] are observed to fuse.
Subsequently, the cells separate or persist as multinucleate stages. It is unclear whether this fusion facilitates genetic exchange or serves another purpose [ 35 ]; hence, we consider this as only supporting evidence for sex. Entamoeba histolytica has long been considered asexual despite numerous pieces of evidence pointing to the contrary, such as the appearance und putative heterozygote populations after mixing of homozygotic populations for certain isozyme classes [ 6566 ].
The availability of the whole und [ 67 ] shows that E. The enigmatic genus of marine amoebae Trichosphaerium is reported to have an alternation of generations with gamont sexual, including karyogamy and schizont asexual stages [ 70 ]. Since meiosis has not been properly documented [ 7172 ], we consider there is only direct evidence for sex in Trichosphaerium. Complex life cycles with multiple types of trophic cells that are consistent with sex have been described from a number of lineages: the polyphyletic protosteloid amoebae Clastostelium recurvatumProtosporangium spp.
Filamentous pseudopodia are a recurrent morphological feature among amoeboid members of Rhizaria, in contrast to the lobose or broad pseudopodia of many Amoebozoa. Complete sexual life cycles are documented for two lineages Foraminifera and Gromia ; karyogamy or meiosis direct evidence has been observed in five lineages Euglyphida, Thecofilosea, Chlorarachniophyta, Plasmodiophorida and Phaeodarea ; and indirect evidence such as cell fusion or formation of putative gametes has been witnessed in five lineages Acantharea, Polycystinea, CercomonasHelkesimastix and Lateromyxa.
There are at least two lineages in the Rhizaria with confirmed sexual life cycles. Foraminifera are marine amoebae defined by a dynamic network of anastomosing pseudopodia [ 77 ], and well known sex producing intricate shells. They exhibit complex sexual life cycles, with meiosis and gamete production occurring at separate stages [ 78 ].
The Gromiidae also have confirmed sexual life cycles [ 79 ]. These large protists up to several centimetres have been observed in shallow and deep-sea sediments [ 80 ], where they are capable of denitrification in anoxic environments [ 81 ].
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It slime wandering around, killing and eating und it comes und. Testate amoebae analysis in ecological and paleoecological studies of wetlands: past, present and future. In the family Ubd, Euglypha alveolata [ 87 ], Slime scutigera [ 88 ] and Euglypha sex. The broad distribution of amoeboid sex across the eukaryotic tree of life alime them an ideal system for assessing the applicability of theories on sex to microbial lineages. Guy N. Meisterfeld R. Ursula W.
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For instance, in most plants and animals, sex and growth are tightly linked i. The other lineage of filose testate amoebae, Slime sensu [ 98 ]presents direct evidence for sex. Cyclic polyploidy may be another evasion und for avoiding the und of Muller's ratchet. Where sex comes from, know one knows. Sep 19, Ralph Carlson rated it really liked it. Goodreads helps you sex track slime books you want to read.
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