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Grevy’s zebra.

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Presentation on theme: "Grevy’s zebra."— Presentation transcript:

1 Grevy’s zebra

2 Grant’s zebra

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4 Grant’s zebra

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6 Speciation Biological species definition: “... groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups.” - Mayr, 1942 One of the YEC claims is that the evolution of organisms can never be experimentally validated because it happened in the past and it takes too long. But in fact we can infer the evolution of organisms by investigating the fossil record. Inference is a perfectly acceptable way of validating a hypothesis. Some branches of biology such as ecology and natural history rely heavily inference. The work of Jane Goodall and Diane Fosey was purely inferential in nature and revolutionized our understanding of chimps and gorillas. Further more, the fact is evolution happened in the past, it’s happening today and it will happen tomorrow. As a consequence we can observe evolution today both in the lab and in nature. One of the experiments we do in my microbiology class is to begin with a bacterium that is originally susceptible to an antibiotic. We then grow up lots and lots of it and we always have one or two of those bacteria that incur a mutation and become resistant to the antibiotic. That is evolution. We also have numerous examples of one species diverging into two different species. Galeopsis tetrahit (hemp-nettle, a type of mint), which was made artificially by hybridizing G. pubescens and G. speciosa - Müntzing, Hereditas, Evening Primrose (Oenothera gigas) While studying the genetics of the evening primrose, Oenothera lamarckiana, de Vries (1905) found an unusual variant among his plants. O. lamarckiana has a chromosome number of 2N = 14. The variant had a chromosome number of 2N = 28. He found that he was unable to breed this variant with O. lamarckiana. He named this new species O. gigas. Kew Primrose (Primula kewensis) Digby (1912) crossed the primrose species Primula verticillata and P. floribunda to produce a sterile hybrid. Polyploidization occurred in a few of these plants to produce fertile offspring. The new species was named P. kewensis. Newton and Pellew (1929) note that spontaneous hybrids of P. verticillata and P. floribunda set tetraploid seed on at least three occasions. These happened in 1905, 1923 and 1926. The Russian cytologist Karpchenko (1927, 1928) crossed the radish, Raphanus sativus, with the cabbage, Brassica oleracea. Despite the fact that the plants were in different genera, he got a sterile hybrid. Some unreduced gametes were formed in the hybrids. This allowed for the production of seed. Plants grown from the seeds were interfertile with each other. They were not interfertile with either parental species. Unfortunately the new plant (genus Raphanobrassica) had the foliage of a radish and the root of a cabbage. 5.5.1 Apple Maggot Fly (Rhagoletis pomonella) Rhagoletis pomonella is a fly that is native to North America. Its normal host is the hawthorn tree. Sometime during the nineteenth century it began to infest apple trees. Since then it has begun to infest cherries, roses, pears and possibly other members of the rosaceae. Quite a bit of work has been done on the differences between flies infesting hawthorn and flies infesting apple. There appear to be differences in host preferences among populations. Offspring of females collected from on of these two hosts are more likely to select that host for oviposition (Prokopy et al. 1988). Genetic differences between flies on these two hosts have been found at 6 out of 13 allozyme loci (Feder et al. 1988, see also McPheron et al. 1988). Laboratory studies have shown an asynchrony in emergence time of adults between these two host races (Smith 1988). Flies from apple trees take about 40 days to mature, whereas flies from hawthorn trees take days to mature. This makes sense when we consider that hawthorn fruit tends to mature later in the season that apples. Hybridization studies show that host preferences are inherited, but give no evidence of barriers to mating. This is a very exciting case. It may represent the early stages of a sympatric speciation event (considering the dispersal of R. pomonella to other plants it may even represent the beginning of an adaptive radiation). It is important to note that some of the leading researchers on this question are urging caution in interpreting it. Feder and Bush (1989) stated: "Hawthorn and apple "host races" of R. pomonella may therefore represent incipient species. However, it remains to be seen whether host-associated traits can evolve into effective enough barriers to gene flow to result eventually in the complete reproductive isolation of R. pomonella populations." 5.5.2 Gall Former Fly (Eurosta solidaginis) Eurosta solidaginis is a gall forming fly that is associated with goldenrod plants. It has two hosts: over most of its range it lays its eggs in Solidago altissima, but in some areas it uses S. gigantea as its host. Recent electrophoretic work has shown that the genetic distances among flies from different sympatric hosts species are greater than the distances among flies on the same host in different geographic areas (Waring et al. 1990). This same study also found reduced variability in flies on S. gigantea. This suggests that some E. solidaginis have recently shifted hosts to this species. A recent study has compared reproductive behavior of the flies associated with the two hosts (Craig et al. 1993). They found that flies associated with S. gigantea emerge earlier in the season than flies associated with S. altissima. In host choice experiments, each fly strain ovipunctured its own host much more frequently than the other host. Craig et al. (1993) also performed several mating experiments. When no host was present and females mated with males from either strain, if males from only one strain were present. When males of both strains were present, statistically significant positive assortative mating was seen. In the presence of a host, assortative mating was also seen. When both hosts and flies from both populations were present, females waited on the buds of the host that they are normally associated with. The males fly to the host to mate. Like the Rhagoletis case above, this may represent the beginning of a sympatric speciation.

7 Beefalo

8 Zebroid (Zorse)

9 Ligers and Tigons, oh my!

10 leopons and liguars

11 Savannah Cat

12 Wholphin

13 Ring Species

14 Schloss, et al, AEM, in press
Displays all OTU at predefined cutoffs User defined OTU algorithms: nearest-neighbor (single-linkage) furthest-neighbor (complete linkage) Average-neighbor (UPGMA)

15 How do you define an Operational Taxanomic Unit (OTU)?
SSU rRNA surveys have traditionally used a 3% or 1% identity cut-off. “…organisms that have less than 97.0% sequence homology will not reassociate to more than 60%…”

16 Polyphyletic traits among bacteria
Brock’s Microbiology of Microorganisms, 14th ed

17 Dissimilarity Matrix and UPGMA tree
E. coli R. rickettsii B. bergdorferi S. cerevisiae H. Sapiens H. marismortui 1.0 0.71 0.64 0.50 0.46 0.67 0.29 0.65 0.36 0.35 0.52 0.6 0.80 0.47 0.48 0.20 0.33 0.40 0.53

18 Speciation Biological species definition: “... groups of actually or potentially interbreeding natural populations which are reproductively isolated from other such groups.” - Mayr, 1942 Speciation in plants: Galeopsis tetrahit, which was made artificially by hybridizing G. pubescens and G. speciosa - Müntzing, 1930 Two species of primrose - deVries, 1905 & Digby, 1912 Raphanobrassica - Karpchenko, 1927 8 more examples Speciation in animals: Apple maggot flies (Rhagoletis pomonella) - Feder et al, 1988 Gall Former flies (Eurosta solidaginis) - Craig et al, 1993 Fruit flies (Drosophila paulistorum) - Dobzhansky & Pavlovsky, 1971 Hybrid from Aspidoscelis exsanguis and Aspidoscelis inornata, Lutes, et al, 2011 One of the YEC claims is that the evolution of organisms can never be experimentally validated because it happened in the past and it takes too long. But in fact we can infer the evolution of organisms by investigating the fossil record. Inference is a perfectly acceptable way of validating a hypothesis. Some branches of biology such as ecology and natural history rely heavily inference. The work of Jane Goodall and Diane Fosey was purely inferential in nature and revolutionized our understanding of chimps and gorillas. Further more, the fact is evolution happened in the past, it’s happening today and it will happen tomorrow. As a consequence we can observe evolution today both in the lab and in nature. One of the experiments we do in my microbiology class is to begin with a bacterium that is originally susceptible to an antibiotic. We then grow up lots and lots of it and we always have one or two of those bacteria that incur a mutation and become resistant to the antibiotic. That is evolution. We also have numerous examples of one species diverging into two different species. Galeopsis tetrahit (hemp-nettle, a type of mint), which was made artificially by hybridizing G. pubescens and G. speciosa - Müntzing, Hereditas, Evening Primrose (Oenothera gigas) While studying the genetics of the evening primrose, Oenothera lamarckiana, de Vries (1905) found an unusual variant among his plants. O. lamarckiana has a chromosome number of 2N = 14. The variant had a chromosome number of 2N = 28. He found that he was unable to breed this variant with O. lamarckiana. He named this new species O. gigas. Kew Primrose (Primula kewensis) Digby (1912) crossed the primrose species Primula verticillata and P. floribunda to produce a sterile hybrid. Polyploidization occurred in a few of these plants to produce fertile offspring. The new species was named P. kewensis. Newton and Pellew (1929) note that spontaneous hybrids of P. verticillata and P. floribunda set tetraploid seed on at least three occasions. These happened in 1905, 1923 and 1926. The Russian cytologist Karpchenko (1927, 1928) crossed the radish, Raphanus sativus, with the cabbage, Brassica oleracea. Despite the fact that the plants were in different genera, he got a sterile hybrid. Some unreduced gametes were formed in the hybrids. This allowed for the production of seed. Plants grown from the seeds were interfertile with each other. They were not interfertile with either parental species. Unfortunately the new plant (genus Raphanobrassica) had the foliage of a radish and the root of a cabbage. 5.5.1 Apple Maggot Fly (Rhagoletis pomonella) Rhagoletis pomonella is a fly that is native to North America. Its normal host is the hawthorn tree. Sometime during the nineteenth century it began to infest apple trees. Since then it has begun to infest cherries, roses, pears and possibly other members of the rosaceae. Quite a bit of work has been done on the differences between flies infesting hawthorn and flies infesting apple. There appear to be differences in host preferences among populations. Offspring of females collected from on of these two hosts are more likely to select that host for oviposition (Prokopy et al. 1988). Genetic differences between flies on these two hosts have been found at 6 out of 13 allozyme loci (Feder et al. 1988, see also McPheron et al. 1988). Laboratory studies have shown an asynchrony in emergence time of adults between these two host races (Smith 1988). Flies from apple trees take about 40 days to mature, whereas flies from hawthorn trees take days to mature. This makes sense when we consider that hawthorn fruit tends to mature later in the season that apples. Hybridization studies show that host preferences are inherited, but give no evidence of barriers to mating. This is a very exciting case. It may represent the early stages of a sympatric speciation event (considering the dispersal of R. pomonella to other plants it may even represent the beginning of an adaptive radiation). It is important to note that some of the leading researchers on this question are urging caution in interpreting it. Feder and Bush (1989) stated: "Hawthorn and apple "host races" of R. pomonella may therefore represent incipient species. However, it remains to be seen whether host-associated traits can evolve into effective enough barriers to gene flow to result eventually in the complete reproductive isolation of R. pomonella populations." 5.5.2 Gall Former Fly (Eurosta solidaginis) Eurosta solidaginis is a gall forming fly that is associated with goldenrod plants. It has two hosts: over most of its range it lays its eggs in Solidago altissima, but in some areas it uses S. gigantea as its host. Recent electrophoretic work has shown that the genetic distances among flies from different sympatric hosts species are greater than the distances among flies on the same host in different geographic areas (Waring et al. 1990). This same study also found reduced variability in flies on S. gigantea. This suggests that some E. solidaginis have recently shifted hosts to this species. A recent study has compared reproductive behavior of the flies associated with the two hosts (Craig et al. 1993). They found that flies associated with S. gigantea emerge earlier in the season than flies associated with S. altissima. In host choice experiments, each fly strain ovipunctured its own host much more frequently than the other host. Craig et al. (1993) also performed several mating experiments. When no host was present and females mated with males from either strain, if males from only one strain were present. When males of both strains were present, statistically significant positive assortative mating was seen. In the presence of a host, assortative mating was also seen. When both hosts and flies from both populations were present, females waited on the buds of the host that they are normally associated with. The males fly to the host to mate. Like the Rhagoletis case above, this may represent the beginning of a sympatric speciation.

19 Apple Maggot & Hawthorn Berry Maggot Flies
Rhagoletis pomonella

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23 Sutter, et al, Science, April 6, 2007

24 QTL & Fixed Mutations in Dogs
Sutter, et al, Science, April 6, 2007

25 SNP 5 in Dogs Sutter, et al, Science, April 6, 2007

26 -Vila, et al. 1997

27 Phylogeny of dogs Lindblad-Toh et al, Nature, 2005
The phylogenetic tree is based on approx15 kb of exon and intron sequence (see text). Branch colours identify the red-fox-like clade (red), the South American clade (green), the wolf-like clade (blue) and the grey and island fox clade (orange). The tree shown was constructed using maximum parsimony as the optimality criterion and is the single most parsimonious tree. Bootstrap values and bayesian posterior probability values are listed above and below the internodes, respectively; dashes indicate bootstrap values below 50% or bayesian posterior probability values below 95%. Horizontal bars indicate indels, with the number of indels shown in parentheses if greater than one. Underlined species names are represented with corresponding illustrations. (Copyright permissions for illustrations are listed in the Supplementary Information.) Divergence time, in millions of years (Myr), is indicated for three nodes as discussed in ref. 1. For scientific names and species descriptions of canids, see ref A tree based on bayesian inference differs from the tree shown in two respects: it groups the raccoon dog and bat-eared fox as sister taxa, and groups the grey fox and island fox as basal to the clade containing these sister taxa. However, neither of these topological differences is strongly supported (see text and Supplementary Information). Lindblad-Toh et al, Nature, 2005

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