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Welcome to the 31st IUBS General Assembly and Conference on ...

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embryos of medaka fish. We found that most of <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

genomic DNA is highly methylated but loci where H3K4<br />

<str<strong>on</strong>g>and</str<strong>on</strong>g>/or H3K27 methylati<strong>on</strong> occur are excepti<strong>on</strong>ally<br />

hypomethylated. Am<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g>m, ~200 loci were found <str<strong>on</strong>g>to</str<strong>on</strong>g><br />

possess l<strong>on</strong>g DNA hypomethylated domain (>5kb) with<br />

bivalent his<str<strong>on</strong>g>to</str<strong>on</strong>g>ne modificati<strong>on</strong>s. Interestingly, ~75 % of<br />

genes of such loci are classified as key developmental<br />

genes including transcripti<strong>on</strong>al regula<str<strong>on</strong>g>to</str<strong>on</strong>g>rs such as<br />

homeobox <str<strong>on</strong>g>and</str<strong>on</strong>g> T‐box genes. The subsequent analysis<br />

using adult liver <str<strong>on</strong>g>and</str<strong>on</strong>g> muscle revealed that those key genes<br />

are subject <str<strong>on</strong>g>to</str<strong>on</strong>g> DNA methylati<strong>on</strong>, except for H3K4me‐rich<br />

promoter regi<strong>on</strong>s, when actively transcribed. Our<br />

time‐course data suggest that decrease in H3K27me<br />

occurs first, prefiguring DNA methylati<strong>on</strong> at later stages.<br />

Toge<str<strong>on</strong>g>the</str<strong>on</strong>g>r with detailed examinati<strong>on</strong> <strong>on</strong> some specific<br />

genes, we propose that key developmental genes are<br />

regulated in a step‐wise manner throughout life, bivalent<br />

<str<strong>on</strong>g>and</str<strong>on</strong>g> DNA hypomethylated state in undifferentiated early<br />

embryos, his<str<strong>on</strong>g>to</str<strong>on</strong>g>ne‐modificati<strong>on</strong>‐dependent inducti<strong>on</strong><br />

during development <str<strong>on</strong>g>and</str<strong>on</strong>g> finally <str<strong>on</strong>g>the</str<strong>on</strong>g> maintenance of<br />

transcripti<strong>on</strong> by DNA‐methylati<strong>on</strong> after birth.<br />

Evoluti<strong>on</strong> as a scientific <str<strong>on</strong>g>the</str<strong>on</strong>g>ory: evidence<br />

from remarkable Chinese fossils<br />

Zh<strong>on</strong>ghe ZHOU<br />

Institute of Vertebrate Pale<strong>on</strong><str<strong>on</strong>g>to</str<strong>on</strong>g>logy <str<strong>on</strong>g>and</str<strong>on</strong>g> Paleoanthropology, CAS,<br />

Beijing, China. Email: Zh<strong>on</strong>ghe@yeah.net;<br />

zhouzh<strong>on</strong>ghe@ivpp.ac.cn<br />

When Darwin wrote On <str<strong>on</strong>g>the</str<strong>on</strong>g> Origin of Species(1859), <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

fossil record was poorly sampled <str<strong>on</strong>g>and</str<strong>on</strong>g> largely incomplete,<br />

yet <str<strong>on</strong>g>the</str<strong>on</strong>g> knowledge of pale<strong>on</strong><str<strong>on</strong>g>to</str<strong>on</strong>g>logy <str<strong>on</strong>g>and</str<strong>on</strong>g> geology cultivated<br />

in <str<strong>on</strong>g>the</str<strong>on</strong>g> nineteen century played a fundamental role in<br />

affirming Darwin’s idea of evoluti<strong>on</strong>. In <str<strong>on</strong>g>the</str<strong>on</strong>g> 1930s–1940s,<br />

Simps<strong>on</strong> brought pale<strong>on</strong><str<strong>on</strong>g>to</str<strong>on</strong>g>logy in<str<strong>on</strong>g>to</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g> evoluti<strong>on</strong>ary<br />

syn<str<strong>on</strong>g>the</str<strong>on</strong>g>sis that c<strong>on</strong>stitutes modern evoluti<strong>on</strong>ary <str<strong>on</strong>g>the</str<strong>on</strong>g>ory.<br />

With str<strong>on</strong>g scientific backing, he established <str<strong>on</strong>g>the</str<strong>on</strong>g> study of<br />

large‐scale evoluti<strong>on</strong>, or macroevoluti<strong>on</strong>, distinct from<br />

evoluti<strong>on</strong> in a general sense (or microevoluti<strong>on</strong>). Ano<str<strong>on</strong>g>the</str<strong>on</strong>g>r<br />

major <str<strong>on</strong>g>the</str<strong>on</strong>g>oretical c<strong>on</strong>tributi<strong>on</strong> by pale<strong>on</strong><str<strong>on</strong>g>to</str<strong>on</strong>g>logists<br />

occurred in <str<strong>on</strong>g>the</str<strong>on</strong>g> 1970s, when Aldridge <str<strong>on</strong>g>and</str<strong>on</strong>g> Gould<br />

proposed <str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>ory of punctuated equilibrium <str<strong>on</strong>g>to</str<strong>on</strong>g><br />

challenge <str<strong>on</strong>g>the</str<strong>on</strong>g> gradualism <str<strong>on</strong>g>the</str<strong>on</strong>g>ory of evoluti<strong>on</strong>.<br />

Fossils provide <str<strong>on</strong>g>the</str<strong>on</strong>g> <strong>on</strong>ly direct evidence of <str<strong>on</strong>g>the</str<strong>on</strong>g> his<str<strong>on</strong>g>to</str<strong>on</strong>g>ry of<br />

evoluti<strong>on</strong>. Fossils that fill <str<strong>on</strong>g>the</str<strong>on</strong>g>oretical evoluti<strong>on</strong>ary<br />

intervals or gaps between existing taxa are often called<br />

‘missing links’. In <str<strong>on</strong>g>the</str<strong>on</strong>g> 1850s, missing links were rare<br />

<str<strong>on</strong>g>and</str<strong>on</strong>g> major gaps existed in <str<strong>on</strong>g>the</str<strong>on</strong>g> fossil record; however<br />

<str<strong>on</strong>g>to</str<strong>on</strong>g>day, more than a century later, this is no l<strong>on</strong>ger <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

case.<br />

Fossil discoveries in <str<strong>on</strong>g>the</str<strong>on</strong>g> past 2 decades, particularly<br />

from China, have greatly enriched our underst<str<strong>on</strong>g>and</str<strong>on</strong>g>ing of<br />

<str<strong>on</strong>g>the</str<strong>on</strong>g> evoluti<strong>on</strong>ary tree of life <str<strong>on</strong>g>and</str<strong>on</strong>g> enhanced <str<strong>on</strong>g>the</str<strong>on</strong>g> evidence<br />

in support of Darwin’s evoluti<strong>on</strong>ary <str<strong>on</strong>g>the</str<strong>on</strong>g>ory. Probably<br />

most remarkable am<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g>se discoveries are <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

earliest vertebrates uncovered from <str<strong>on</strong>g>the</str<strong>on</strong>g> Early Cambrian,<br />

<str<strong>on</strong>g>the</str<strong>on</strong>g> oldest <str<strong>on</strong>g>and</str<strong>on</strong>g> most primitive turtle from <str<strong>on</strong>g>the</str<strong>on</strong>g> Middle<br />

Triassic, <str<strong>on</strong>g>and</str<strong>on</strong>g> four‐winged dinosaurs that are am<strong>on</strong>g <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

closest relatives <str<strong>on</strong>g>to</str<strong>on</strong>g> birds. Fur<str<strong>on</strong>g>the</str<strong>on</strong>g>rmore, studies <strong>on</strong><br />

Chinese fossils have shed new light <strong>on</strong> major patterns of<br />

biological radiati<strong>on</strong> <str<strong>on</strong>g>and</str<strong>on</strong>g> extincti<strong>on</strong> events during <str<strong>on</strong>g>the</str<strong>on</strong>g> last<br />

500 milli<strong>on</strong> years.<br />

The sudden appearance of advanced animals in <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

Cambrian fossil record, known as ‘Darwin’s headache’,<br />

can now be addressed through discoveries from various<br />

early Cambrian <str<strong>on</strong>g>and</str<strong>on</strong>g> Precambrian lagerstätte; <str<strong>on</strong>g>the</str<strong>on</strong>g><br />

discovery of numerous flowered plants from <str<strong>on</strong>g>the</str<strong>on</strong>g> Early<br />

Cretaceous also helps <str<strong>on</strong>g>to</str<strong>on</strong>g> explain Darwin’s ‘abominable<br />

mystery’. Yet probably <str<strong>on</strong>g>the</str<strong>on</strong>g> most significant of <str<strong>on</strong>g>the</str<strong>on</strong>g>se<br />

Chinese discoveries is <str<strong>on</strong>g>the</str<strong>on</strong>g> discovery of 160 milli<strong>on</strong> year<br />

old fea<str<strong>on</strong>g>the</str<strong>on</strong>g>red dinosaurs that c<strong>on</strong>stitute perfect missing<br />

links between birds <str<strong>on</strong>g>and</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>ir dinosaurian ances<str<strong>on</strong>g>to</str<strong>on</strong>g>rs.<br />

One of <str<strong>on</strong>g>the</str<strong>on</strong>g>se important taxa is named Anchiornis huxleyi,<br />

with its species name in h<strong>on</strong>or of Huxley, Darwin’<br />

bulldog, who was <str<strong>on</strong>g>the</str<strong>on</strong>g> first <str<strong>on</strong>g>to</str<strong>on</strong>g> propose <str<strong>on</strong>g>the</str<strong>on</strong>g> hypo<str<strong>on</strong>g>the</str<strong>on</strong>g>sis of<br />

a dinosaurian origin of birds.<br />

Evoluti<strong>on</strong> as a scientific <str<strong>on</strong>g>the</str<strong>on</strong>g>ory or fact, has withs<str<strong>on</strong>g>to</str<strong>on</strong>g>od<br />

various challenges over <str<strong>on</strong>g>the</str<strong>on</strong>g> past 150 years; however,<br />

<str<strong>on</strong>g>the</str<strong>on</strong>g> <str<strong>on</strong>g>the</str<strong>on</strong>g>ory itself is still evolving <str<strong>on</strong>g>and</str<strong>on</strong>g> probably can never<br />

be called complete. While pale<strong>on</strong><str<strong>on</strong>g>to</str<strong>on</strong>g>logy c<strong>on</strong>tinues <str<strong>on</strong>g>to</str<strong>on</strong>g><br />

c<strong>on</strong>tribute evidence of <str<strong>on</strong>g>the</str<strong>on</strong>g> fact of evoluti<strong>on</strong> <str<strong>on</strong>g>and</str<strong>on</strong>g> elucidate<br />

details of evoluti<strong>on</strong>ary his<str<strong>on</strong>g>to</str<strong>on</strong>g>ry, we remain a l<strong>on</strong>g way<br />

from filling <str<strong>on</strong>g>the</str<strong>on</strong>g> gap between our underst<str<strong>on</strong>g>and</str<strong>on</strong>g>ing of<br />

evoluti<strong>on</strong> <strong>on</strong> <str<strong>on</strong>g>the</str<strong>on</strong>g> micro‐ <str<strong>on</strong>g>and</str<strong>on</strong>g> macroevoluti<strong>on</strong>ary scale.<br />

Evolvability in sequence space<br />

Daniel L HARTL<br />

Department of Organismic <str<strong>on</strong>g>and</str<strong>on</strong>g> Evoluti<strong>on</strong>ary Biology, Harvard<br />

University, Cambridge, MA 02138, USA. Email:<br />

dhartl@oeb.harvard.edu<br />

49

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