ࡱ> b00000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000000003 )jbjb^^ Zh<h<B(l8r|uc^* "BBB2O [[[[[[[[,d f[ccc[3BB*333clBBU 3&DjDc[337 RlT<5W/ xS*TucucSJg3gT3Differentiation Waves vs Positional Information, Genome as a Machine vs Gene Networks: Which Paradigms will Prevail? Richard Gordon Departments of Radiology and Electrical & Computer Engineering University of Manitoba, Room GA216, Health Sciences Centre 820 Sherbrook Street, Winnipeg, MB R3A 1R9 Canada Phone: (204) 789-3828, Fax: (204) 787-2080, E-mail: GordonR@ms.UManitoba.ca Invited August, 2000 by: Dr. Adam S. Wilkins, Editor Editorial Office, BioEssays, Units10/11, Tredgold Lane, Napier Street Cambridge CB1 1HN, United Kingdom Phone: +44 (1223) 357542, Fax: +44 (1223) 359761 E-mail: edoffice@bioessays.demon.co.uk Prepared for BioEssays. Manuscript started August 8, 2000. Version 32, submitted December 10, 2001, compressed Version 41, January 1, 2002. Abstract Positional information via gradients of morphogens has been the dominant paradigm in developmental biology for the past three decades, but is beginning to show strains. The major alternative paradigm for cells in development is that a differentiation wave is the trigger for each change in gene expression. These waves, including the morphogenetic furrow of the Drosophila eye imaginal disc, have hitherto been dismissed as epiphenomena. Cell differentiation is paralleled by unknown events in the nucleus. Here the paradigm of basins of attraction in networks of regulatory genes and proteins has held sway equally long. Alternatively the genome is a machine for differentiation, with discrete physical states for each cell type that switch the cell between networks. This can be modeled by toy Wurfels. Practical consequences include a suggested tissue synthesizer that could turn stem cells into any desired cell type. We need to choose which pair of cellular and nuclear paradigms is correct. Keywords: differentiation waves, positional information, genome as machine, gene networks, paradigms, stem cells, embryonic induction Introduction Both Morgan  QUOTE "{Morgan 1901 #364770}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Morgan 1901 #364770}\00\15\00{^\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Morgan 1901 #364770\00\13\00 {Morgan 1901 #364770} and Nieuwkoop  QUOTE "{Nieuwkoop 1973 #380920}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Nieuwkoop 1973 #380920}\00\18\00\07\11\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Nieuwkoop 1973 #380920\00\16\00 {Nieuwkoop 1973 #380920} held dim views of gradients as explanations of differentiation during embryogenesis. With Wolperts introduction of the concept of positional information  QUOTE "{Wolpert 1969 #572290}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{Wolpert 1969 #572290}\00\16\00A\17\00\00%Dick 9:Dick's Databases:Dick's Biblio\14Wolpert 1969 #572290\00\14\00 {Wolpert 1969 #572290}, gradients of positional information have nevertheless become the predominant paradigm of our time. In the major alternative paradigm, cell differentiation is triggered by waves of cell contraction or expansion, and gradients are epiphenomena  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660}. Little attention has been paid to mechanisms in the nucleus that correspond to cell differentiation in intact embryos. Driven by the wealth of data from genome sequencing and proteomics, some researchers assume that differentiation involves vast arrays of interacting genes and gene products. The basic idea is that genetic networks have multiple stable modes of operation (basins of attraction in the language of dynamics), and that each one corresponds to a state of cell differentiation  QUOTE "{Delbrck 1949a #118650}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Delbrck 1949a #118650}\00\18\00S\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Delbrck 1949a #118650\00\16\00 {Delbrck 1949a #118650}  QUOTE "{Kauffman 1993 #269200}"  ADDIN PROCITE \11\05\19\02\00\00\00\17{Kauffman 1993 #269200}\00\17\00v5\00\00%Dick 9:Dick's Databases:Dick's Biblio\15Kauffman 1993 #269200\00\15\00 {Kauffman 1993 #269200}. An entirely different paradigm is being formulated, in which the cell nucleus has a discrete structure for each cell type, accessible to one another by machine-like alterations of the whole nucleus. These configurational changes would switch the nucleus from one genetic network to another. These four paradigms are intimately linked, because the embryo is first and foremost an array of different kinds of cells from the same clone. The pair of paradigms chosen must simultaneously explain both cellular and nuclear differentiation (Table 1), and must be mechanistically linked together. Nuclear( Cellular(Positional InformationDifferentiation WavesGenetic NetworksGradients & NetworksWaves & NetworksGenome as MachineGradients & WurfelsWaves & WurfelsTable 1. There are four pairs of paradigms between which we may have to choose, to explain the cellular and nuclear bases of embryogenesis. As we shall see below, Wurfels may be taken as working models for the genome as a machine. Gradients in Trouble In computer jargon positional information is equivalent to: 1. a cell knows where it is (which is the role of the gradient); 2. its coordinates are somehow mapped into a set of instructions, as in a look-up table, case statement, etc.; 3. the cell carries out these instructions. Simple gradient models for positional information  QUOTE "{Crick 1970 #106650}"  ADDIN PROCITE \11\05\19\02\00\00\00\14{Crick 1970 #106650}\00\14\00 \05\00\00%Dick 9:Dick's Databases:Dick's Biblio\12Crick 1970 #106650\00\12\00 {Crick 1970 #106650} have been replaced by models requiring co-factors and cooperativity  QUOTE "{Ma, Yuan, et al. 1996 #324510}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Ma, Yuan, et al. 1996 #324510}\00\1F\00%\00\00%Dick 9:Dick's Databases:Dick's Biblio\1DMa, Yuan, et al. 1996 #324510\00\1D\00 {Ma, Yuan, et al. 1996 #324510}  QUOTE "{Almirantis & Papageorgiou 1999 #9180}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Almirantis & Papageorgiou 1999 #9180}\00&\00{\00\00%Dick 9:Dick's Databases:Dick's Biblio$Almirantis & Papageorgiou 1999 #9180\00$\00 {Almirantis & Papageorgiou 1999 #9180}  QUOTE "{Kerszberg 1999 #273830}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Kerszberg 1999 #273830}\00\18\00\15&\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Kerszberg 1999 #273830\00\16\00 {Kerszberg 1999 #273830} and interaction with other purported morphogens  QUOTE "{Podos & Ferguson 1999 #413820}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Podos & Ferguson 1999 #413820}\00\1F\00_0 \00\00%Dick 9:Dick's Databases:Dick's Biblio\1DPodos & Ferguson 1999 #413820\00\1D\00 {Podos & Ferguson 1999 #413820}. Some models  QUOTE "{Lacalli & Harrison 1991 #294770}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Lacalli & Harrison 1991 #294770}\00!\00\16\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FLacalli & Harrison 1991 #294770\00\1F\00 {Lacalli & Harrison 1991 #294770} use a combination of a monotonic gradient of one morphogen with an additional pair of reaction-diffusion  QUOTE "{Turing 1952 #531890}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Turing 1952 #531890}\00\15\00 \15\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Turing 1952 #531890\00\13\00 {Turing 1952 #531890} morphogens. Due to fluctuations in small numbers of morphogen molecules, a gradient sharpening reaction-diffusion mechanism has been further added  QUOTE "{Holloway & Harrison 1999a #608200}"  ADDIN PROCITE \11\05\19\02\00\00\00#{Holloway & Harrison 1999a #608200}\01\05\00\01\00\00\00\00\00\00\0Ae \00\00\00\01\0C{ <\0C{\1F0(\00\04\08 \00\00\00\00\0C{\1F\08\00\10\04\00\00\00\00\00"\0B+"\0C\03\0B+" \0C\03! \0C\03`\0C\03`\0Cx\03\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B`"PH\0Bwa\08\00\10\04\00\00\00\00\00"\0B+"\0C\03\0B+" \0C\03! \0C\03`\0C\03`\0Cx\0F\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B`"PH\0Bwa\1C\00\00%Dick 9:Dick's Databases:Dick's Biblio!Holloway & Harrison 1999a #608200\01\04\00\08\00\10\04\00\00\00\00\00"\0B+"\0C\03\0B+" \0C\03! \0C\03`\0C\03`\0Cx\03\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BI! \0B*0\08\00\10\04\00\00\00\00\00"\0B+"\0C\03\0B+" \0C\03! \0C\03`\0C\03`\0Cx\0E\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BI! \0B*0 {Holloway & Harrison 1999a #608200}. New models for morphogen gradients continue to appear  QUOTE "{Pages & Kerridge 2000 #396880}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Pages & Kerridge 2000 #396880}\00\1F\00]\00\00%Dick 9:Dick's Databases:Dick's Biblio\1DPages & Kerridge 2000 #396880\00\1D\00 {Pages & Kerridge 2000 #396880}, suggesting that our understanding of positional information via gradients is far from complete. Some authors have raised doubt about the applicability of the concept of morphogen gradients  QUOTE "{Reilly & Melton 1996 #431380}"  ADDIN PROCITE \11\05\19\02\00\00\00\1E{Reilly & Melton 1996 #431380}\00\1E\00J\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1CReilly & Melton 1996 #431380\00\1C\00 {Reilly & Melton 1996 #431380}  QUOTE "{Neumann & Cohen 1997 #378300}"  ADDIN PROCITE \11\05\19\02\00\00\00\1E{Neumann & Cohen 1997 #378300}\00\1E\004\00\00%Dick 9:Dick's Databases:Dick's Biblio\1CNeumann & Cohen 1997 #378300\00\1C\00 {Neumann & Cohen 1997 #378300} or positional information  QUOTE "{Lintilhac 1984 #315070}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Lintilhac 1984 #315070}\00\18\00G\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Lintilhac 1984 #315070\00\16\00 {Lintilhac 1984 #315070} to their favorite organisms or body parts. There is a theoretical objection to simple gradients, since receptors for a morphogen would adsorb it, preventing a long range gradient from forming  QUOTE "{Kerszberg & Wolpert 1998a #273880}"  ADDIN PROCITE \11\05\19\02\00\00\00#{Kerszberg & Wolpert 1998a #273880}\01\05\00\01\00\00\00\00\00\00\0Ae \00\00\00\01\0C{ <\0C{\1F0(\00\04\08 \00\00\00\00\0C{\1F\09\00\10\04\00\00\00\00\00"\0B+"\0C\15\08\0B+" \0C\15\0C\15(\0C\15(\0Cx\03\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B`"s"\0Bwa\07\00\10\04\00\00\00\00\00"\0B+"\0C\15\08\0B+" \0C\15\0C\15(\0C\15(\0Cx\0F\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B`"s"\0Bwa \00\00%Dick 9:Dick's Databases:Dick's Biblio!Kerszberg & Wolpert 1998a #273880\01\04\00\09\00\10\04\00\00\00\00\00"\0B+"\0C\15\08\0B+" \0C\15\0C\15(\0C\15(\0Cx\03\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BJ\0B*0\07\00\10\04\00\00\00\00\00"\0B+"\0C\15\08\0B+" \0C\15\0C\15(\0C\15(\0Cx\0E\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BJ\0B*0 {Kerszberg & Wolpert 1998a #273880}. On the other hand, cytonemes  QUOTE "{Ramirez-Weber & Kornberg 1999 #424800}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Ramirez-Weber & Kornberg 1999 #424800}\00'\00/\1B\00\00%Dick 9:Dick's Databases:Dick's Biblio%Ramirez-Weber & Kornberg 1999 #424800\00%\00 {Ramirez-Weber & Kornberg 1999 #424800}, cell extensions 0.2m wide and up to 800m long, have been put forth as potential long range carriers of gradients, though they would presumably only affect the cells onto which they impinge. Turing patterns are sensitive to the shape of the tissue, i.e., not robust for morphogenesis  QUOTE "{Bunow, Kernevez, et al. 1980 #68180}"  ADDIN PROCITE \11\05\19\02\00\00\00%{Bunow, Kernevez, et al. 1980 #68180}\00%\00 )\00\00%Dick 9:Dick's Databases:Dick's Biblio#Bunow, Kernevez, et al. 1980 #68180\00#\00 {Bunow, Kernevez, et al. 1980 #68180}, and may prove counterproductive additions to gradient models. One criterion for a model of embryogenesis is universality across species, because embryos are related through evolution. For example, we cannot regard the maternal bicoid gradient, found in Drosophila, as universal because: Any maternal gradients are probably lost during clone formation of thousands of eggs from one in polyembryonic wasps  QUOTE "{Grbic, Nagy, et al. 1998 #194360}"  ADDIN PROCITE \11\05\19\02\00\00\00"{Grbic, Nagy, et al. 1998 #194360}\00"\00" H"\00\00%Dick 9:Dick's Databases:Dick's Biblio Grbic, Nagy, et al. 1998 #194360\00 \00 {Grbic, Nagy, et al. 1998 #194360}. Short germ band insects, such as the flour beetle Tribolium, do not form a syncytial blastoderm  QUOTE "{Schaeffer, Killian, et al. 2000 #614220}"  ADDIN PROCITE \11\05\19\02\00\00\00){Schaeffer, Killian, et al. 2000 #614220}\00)\00`\00\00%Dick 9:Dick's Databases:Dick's Biblio'Schaeffer, Killian, et al. 2000 #614220\00'\00 {Schaeffer, Killian, et al. 2000 #614220}. The role of bicoid has changed during evolution  QUOTE "{Stauber, Taubert, et al. 2000 #614210}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Stauber, Taubert, et al. 2000 #614210}\00'\00_\00\00%Dick 9:Dick's Databases:Dick's Biblio%Stauber, Taubert, et al. 2000 #614210\00%\00 {Stauber, Taubert, et al. 2000 #614210}. Further back in evolution the developmental importance of bicoid is questionable: as bicoid seems to be a recently acquired gene in flies  QUOTE "{Wimmer, Carleton, et al. 2000 #614250}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Wimmer, Carleton, et al. 2000 #614250}\00'\00c\00\00%Dick 9:Dick's Databases:Dick's Biblio%Wimmer, Carleton, et al. 2000 #614250\00%\00 {Wimmer, Carleton, et al. 2000 #614250}. The first two objections apply to any purported morphogen in early insect embryogenesis. Furthermore, if positional information does not generate gradients, but merely uses them, then it requires a theory of how, where and when gradients are formed in the first place. This is a fundamental obstacle for the theory, because some eggs (the marine alga Fucus  QUOTE "{Jaffe 1968 #252600}"  ADDIN PROCITE \11\05\19\02\00\00\00\14{Jaffe 1968 #252600}\00\14\00/2\00\00%Dick 9:Dick's Databases:Dick's Biblio\12Jaffe 1968 #252600\00\12\00 {Jaffe 1968 #252600}; mammals  QUOTE "{Evsikov, Morozova, et al. 1994 #146230}"  ADDIN PROCITE \11\05\19\02\00\00\00({Evsikov, Morozova, et al. 1994 #146230}\00(\00X\00\00%Dick 9:Dick's Databases:Dick's Biblio&Evsikov, Morozova, et al. 1994 #146230\00&\00 {Evsikov, Morozova, et al. 1994 #146230}) are initially completely spherically symmetric as far as we can tell. We cannot rely on gradients of maternal determinants even in the chick: The chick, for example, may be cut into many fragments at a stage when the embryo has more than 105 cells, and each fragment will spontaneously establish its own polarity and initiate axis development. In these animals, the localization of maternal determinants is unlikely to play a crucial, instructive role  QUOTE "{Perrimon & Stern 1999 #406630}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Perrimon & Stern 1999 #406630}\00\1F\00b\1A\00\00%Dick 9:Dick's Databases:Dick's Biblio\1DPerrimon & Stern 1999 #406630\00\1D\00 {Perrimon & Stern 1999 #406630}. Differentiation Waves as an Alternative to Gradients Although it appears that cascades of gene activation are implicated in the building of many tissues and organs, there cannot be an infinite regression of gene activation by products of genes that must in turn be expressed. At various points in the causal chain of epigenesis, physical factors and nongenetic chemical factors& . may be the early initiators of gene function  QUOTE "{Bell 2000 #644240}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Bell 2000 #644240}\00\13\00S\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Bell 2000 #644240\00\11\00 {Bell 2000 #644240}. It is generally thought that Child  QUOTE "{Child 1914 #89180}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Child 1914 #89180}\00\13\00)\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Child 1914 #89180\00\11\00 {Child 1914 #89180} emphasized gradients, albeit of metabolic properties. However, he actually supported the alternative wave paradigm, suggesting that gradients are consequences of these waves, not vice versa: ...the influence of a dominant region is primarily transmissive, rather than transportive in character, that is, that it consists in some form of energy transmission in protoplasm or along limiting surfaces, rather than in mass transport of substance  QUOTE "{Child 1928 #89230}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Child 1928 #89230}\00\13\00H#\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Child 1928 #89230\00\11\00 {Child 1928 #89230}. This is the basic idea of differentiation waves. In fact, it provides a test for distinguishing morphogen gradient models  QUOTE "{Tabata 2001 #1570}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Tabata 2001 #1570}\00\13\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Tabata 2001 #1570\00\11\00 {Tabata 2001 #1570} (Figure1a,b,c) from differentiation waves (Figure1d), perhaps by radioactive labeling or fluorescent tagging of purported morphogens. The mere presence of a gradient of a purported morphogen does not tell us if the gradient is cause or effect. Child  QUOTE "{Child 1928 #89230}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Child 1928 #89230}\00\13\00H#\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Child 1928 #89230\00\11\00 {Child 1928 #89230} was preceded by vonUexkll  QUOTE "{vonUexkll 1926 #546050}"  ADDIN PROCITE \11\05\19\02\00\00\00\1A{vonUexkll 1926 #546050}\00\1A\00n\1B\00\00%Dick 9:Dick's Databases:Dick's Biblio\18vonUexkll 1926 #546050\00\18\00 {vonUexkll 1926 #546050}, who suggested nonmaterial impulses that not only are transmitted from cell to cell, but also trigger specific gene expression. The idea that waves are involved in embryogenesis goes back at least to 1888  QUOTE "{Whitman 1888 #563470}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{Whitman 1888 #563470}\00\16\00AQ\00\00%Dick 9:Dick's Databases:Dick's Biblio\14Whitman 1888 #563470\00\14\00 {Whitman 1888 #563470}. Differentiation waves  QUOTE "{Gordon & Brodland 1987a #188950}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Gordon & Brodland 1987a #188950}\00!\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FGordon & Brodland 1987a #188950\00\1F\00 {Gordon & Brodland 1987a #188950}  QUOTE "{Gordon, Bjrklund, et al. 1994 #188900}"  ADDIN PROCITE \11\05\19\02\00\00\00({Gordon, Bjrklund, et al. 1994 #188900}\00(\00V%\00\00%Dick 9:Dick's Databases:Dick's Biblio&Gordon, Bjrklund, et al. 1994 #188900\00&\00 {Gordon, Bjrklund, et al. 1994 #188900} are a set of slow to ultraslow  QUOTE "{Jaffe 1999 #252760}"  ADDIN PROCITE \11\05\19\02\00\00\00\14{Jaffe 1999 #252760}\00\14\00\18\16\00\00%Dick 9:Dick's Databases:Dick's Biblio\12Jaffe 1999 #252760\00\12\00 {Jaffe 1999 #252760} physical expansion and contraction waves that correlate spatially and temporally with the formation of tissues in the early urodele amphibian embryo. If this correlation is causal, and if we could explain these waves and how, when and where they are triggered, then we might be well on the way to a complete solution to embryogenesis. The ectoderm contraction wave (Figure2) leaves the neural plate (neuroepithelium) in its wake  QUOTE "{Brodland, Gordon, et al. 1994 #62320}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Brodland, Gordon, et al. 1994 #62320}\00&\00!\00\00%Dick 9:Dick's Databases:Dick's Biblio$Brodland, Gordon, et al. 1994 #62320\00$\00 {Brodland, Gordon, et al. 1994 #62320} in the urodele amphibian Ambystoma mexicanum (the axolotl). A similar wave may be awaiting discovery beneath the superficial epithelium of the anuran amphibian Xenopus  QUOTE "{Nieuwkoop, Bjrklund, et al. 1996 #381050}"  ADDIN PROCITE \11\05\19\02\00\00\00+{Nieuwkoop, Bjrklund, et al. 1996 #381050}\00+\00D\00\00%Dick 9:Dick's Databases:Dick's Biblio)Nieuwkoop, Bjrklund, et al. 1996 #381050\00)\00 {Nieuwkoop, Bjrklund, et al. 1996 #381050}. Its calcium component has probably already been seen  QUOTE "{Leclerc, Webb, et al. 2000 #636020}"  ADDIN PROCITE \11\05\19\02\00\00\00${Leclerc, Webb, et al. 2000 #636020}\00$\00p\00\00%Dick 9:Dick's Databases:Dick's Biblio"Leclerc, Webb, et al. 2000 #636020\00"\00 {Leclerc, Webb, et al. 2000 #636020}  QUOTE "{Wallingford, Ewald, et al. 2001 #666520}"  ADDIN PROCITE \11\05\19\02\00\00\00){Wallingford, Ewald, et al. 2001 #666520}\01\05\00\01\00\00\00\00\00\00\0Ae \00\00\00\01\0C{ <\0C{\1F0(\00\04\08 \00\00\00\00\0C{\1F\0B\00\10\04\00\00\00\00\00"\0E.~\0Eq\0E.(\0Eq& \0EqU\0EqU\0F4k\02\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B @\0Bwa\05\00\10\04\00\00\00\00\00"\0E.~\0Eq\0E.(\0Eq& \0EqU\0EqU\0F4k\16\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B @\0Bwa\06\01\00%Dick 9:Dick's Databases:Dick's Biblio'Wallingford, Ewald, et al. 2001 #666520\01\04\00\0B\00\10\04\00\00\00\00\00"\0E.~\0Eq\0E.(\0Eq& \0EqU\0EqU\0F4k\02\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BJh\0B*0\05\00\10\04\00\00\00\00\00"\0E.~\0Eq\0E.(\0Eq& \0EqU\0EqU\0F4k\15\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BJh\0B*0 {Wallingford, Ewald, et al. 2001 #666520}. We have used a sequence of contraction and expansion waves  QUOTE "{Bjrklund & Gordon 1994 #45690}"  ADDIN PROCITE \11\05\19\02\00\00\00 {Bjrklund & Gordon 1994 #45690}\00 \00O<\00\00%Dick 9:Dick's Databases:Dick's Biblio\1EBjrklund & Gordon 1994 #45690\00\1E\00 {Bjrklund & Gordon 1994 #45690} to explain the shape of the urodele fate map  QUOTE "{Vogt 1929a #544800}"  ADDIN PROCITE \11\05\19\02\00\00\00\14{Vogt 1929a #544800}\00\14\008\16\00\00%Dick 9:Dick's Databases:Dick's Biblio\12Vogt 1929a #544800\00\12\00 {Vogt 1929a #544800}. Contraction waves are much like the morphogenetic furrow in the Drosophila eye imaginal disc  QUOTE "{Poodry, Hall, et al. 1973 #415330}"  ADDIN PROCITE \11\05\19\02\00\00\00#{Poodry, Hall, et al. 1973 #415330}\00#\00\00\00%Dick 9:Dick's Databases:Dick's Biblio!Poodry, Hall, et al. 1973 #415330\00!\00 {Poodry, Hall, et al. 1973 #415330}  QUOTE "{Ready, Hanson, et al. 1976 #429330}"  ADDIN PROCITE \11\05\19\02\00\00\00${Ready, Hanson, et al. 1976 #429330}\00$\00\12\00\00%Dick 9:Dick's Databases:Dick's Biblio"Ready, Hanson, et al. 1976 #429330\00"\00 {Ready, Hanson, et al. 1976 #429330}, except that only a single step of differentiation appears to occur, such as ectoderm to neuroepithelium, rather than multiple steps (forming the 32 cell ommatidia). Expansion waves resemble epiboly, such as the one correlated with differentiation of ectoderm to epidermis  QUOTE "{Gordon, Bjrklund, et al. 1994 #188900}"  ADDIN PROCITE \11\05\19\02\00\00\00({Gordon, Bjrklund, et al. 1994 #188900}\00(\00V%\00\00%Dick 9:Dick's Databases:Dick's Biblio&Gordon, Bjrklund, et al. 1994 #188900\00&\00 {Gordon, Bjrklund, et al. 1994 #188900}. This suggests that each tissue undergoes a binary step of differentiation caused by a contraction/expansion pair of spatially complementary waves, and that embryogenesis can be represented by a simple binary differentiation tree (Figure3). As such, the differentiation tree is the logic of the genome. It depicts how the genetic program plays itself out during development, as a bifurcating alternation of physics and genetics, and forms the fundamental germ-line replicator  QUOTE "{Dawkins 1982 #115370}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{Dawkins 1982 #115370}\00\16\00D\06\00\00%Dick 9:Dick's Databases:Dick's Biblio\14Dawkins 1982 #115370\00\14\00 {Dawkins 1982 #115370} for the evolution of development  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660}. Because differentiation waves are slow, 3m/min in urodeles at 20oC (Figure 2), they should create gradients in a tissue as they traverse it (Figure1d). Each gene product may be turned on or off (or left as is) in a cell as a wave enters it and the cell begins to participate in propagation of the wave. The profile of the gradient behind a wave will depend on the dynamics of synthesis and degradation or reaction of the specific gene product, but one strong prediction is that all gradients will follow waves. The Drosophila morphogenetic furrow shows gradients within and behind it  QUOTE "{Kimmel, Heberlein, et al. 1990 #276670}"  ADDIN PROCITE \11\05\19\02\00\00\00({Kimmel, Heberlein, et al. 1990 #276670}\01\05\00\01\00\00\00\00\00\00\0Ae \00\00\00\01\0C{ <\0C{\1F0(\00\04\08 \00\00\00\00\0C{\1F\06\00\10\04\00\00\00\00\00"\0D\10\0E!\10\0D`"H\0E!u\00\0E!G`\0E!G`\0Eo\02\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B \0Bwa\09\00\10\04\00\00\00\00\00"\0D\10\0E!\10\0D`"H\0E!u\00\0E!G`\0E!G`\0Eo\16\00\00\00\00\00\00{\1C<\0C{\1C<\0C{\1C\0C{\1C\0BW_\0Bw \0B \0Bwa\1FH\00\00%Dick 9:Dick's Databases:Dick's Biblio&Kimmel, Heberlein, et al. 1990 #276670\01\04\00\06\00\10\04\00\00\00\00\00"\0D\10\0E!\10\0D`"H\0E!u\00\0E!G`\0E!G`\0Eo\02\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BI 8\0B*0\09\00\10\04\00\00\00\00\00"\0D\10\0E!\10\0D`"H\0E!u\00\0E!G`\0E!G`\0Eo\15\00\00\00\00\00\00.N\0C.N\0C.O\04\0C.OD\0B\0Ah\0B*d"\10\0BI 8\0B*0 {Kimmel, Heberlein, et al. 1990 #276670}. Purported morphogens ahead of a furrow, such as hedgehog, may be located precisely where contraction starts  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s \00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660}. The furrow only indicates which cells are presently at maximal contraction. Differentiation waves may include the mitotic waves in the Drosophila blastoderm and its later cellular mitotic waves  QUOTE "{Foe 1989 #157560}"  ADDIN PROCITE \11\05\19\02\00\00\00\12{Foe 1989 #157560}\00\12\00+H\00\00%Dick 9:Dick's Databases:Dick's Biblio\10Foe 1989 #157560\00\10\00 {Foe 1989 #157560}, and formation of the sunflower head  QUOTE "{Hernndez & Green 1993 #223840}"  ADDIN PROCITE \11\05\19\02\00\00\00 {Hernndez & Green 1993 #223840}\00 \00\04Q\00\00%Dick 9:Dick's Databases:Dick's Biblio\1EHernndez & Green 1993 #223840\00\1E\00 {Hernndez & Green 1993 #223840}. Two Contesting Paradigms for the Nucleus in Differentiation Is the differentiated state just a particular mode of a vast array of interacting chemical reactions  QUOTE "{Salazar-Ciudad, Garcia-Fernandez, et al. 2000 #596730}"  ADDIN PROCITE \11\05\19\02\00\00\007{Salazar-Ciudad, Garcia-Fernandez, et al. 2000 #596730}\007\00\00\00%Dick 9:Dick's Databases:Dick's Biblio5Salazar-Ciudad, Garcia-Fernandez, et al. 2000 #596730\005\00 {Salazar-Ciudad, Garcia-Fernandez, et al. 2000 #596730}, with the nucleus, like the cytoplasm before it, is treated as a well stirred bag of reactants? Does this model miss the essence of the problem? Is the nuclear structure paramount? Is the nucleus a machine for differentiation? Does it control the network? What is its relationship to morphogen gradients or differentiation waves? Large scale databases are being constructed of interacting proteins  QUOTE "{Xenarios, Fernandez, et al. 2001 #598770}"  ADDIN PROCITE \11\05\19\02\00\00\00*{Xenarios, Fernandez, et al. 2001 #598770}\00*\00Q\00\00%Dick 9:Dick's Databases:Dick's Biblio(Xenarios, Fernandez, et al. 2001 #598770\00(\00 {Xenarios, Fernandez, et al. 2001 #598770} and massive computer simulations of interacting genes and gene products are underway  QUOTE "{Hasty, McMillen, et al. 2001a #641650}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Hasty, McMillen, et al. 2001a #641650}\00'\00q\00\00%Dick 9:Dick's Databases:Dick's Biblio%Hasty, McMillen, et al. 2001a #641650\00%\00 {Hasty, McMillen, et al. 2001a #641650}, with the presumption that the network model is correct. In the network paradigm, the presumption is that the essence of differentiation is a basin of attraction in the high dimensional space of reaction parameters of a huge network of chemical reactions, with a cell settling into one of multiple steady states  QUOTE "{Delbrck 1949a #118650}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Delbrck 1949a #118650}\00\18\00S\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Delbrck 1949a #118650\00\16\00 {Delbrck 1949a #118650}  QUOTE "{Kauffman 1993 #269200}"  ADDIN PROCITE \11\05\19\02\00\00\00\17{Kauffman 1993 #269200}\00\17\00v5\00\00%Dick 9:Dick's Databases:Dick's Biblio\15Kauffman 1993 #269200\00\15\00 {Kauffman 1993 #269200}. For network theory flipping of pathways  QUOTE "{Kauffman 1973 #269070}"  ADDIN PROCITE \11\05\19\02\00\00\00\17{Kauffman 1973 #269070}\00\17\00%\0C\00\00%Dick 9:Dick's Databases:Dick's Biblio\15Kauffman 1973 #269070\00\15\00 {Kauffman 1973 #269070} might be explicable in terms of their organization, but why identical cells should flip different ways to produce the shapes and patterns of an organism has not yet been explained. The spatial problem is not answered by attempts to model differentiation as positive feedbacks between the networks of neighboring cells  QUOTE "{Kaneko & Yomo 1999 #265800}"  ADDIN PROCITE \11\05\19\02\00\00\00\1C{Kaneko & Yomo 1999 #265800}\00\1C\00\1F\00\00%Dick 9:Dick's Databases:Dick's Biblio\1AKaneko & Yomo 1999 #265800\00\1A\00 {Kaneko & Yomo 1999 #265800}  QUOTE "{Huang & Ingber 2000 #594210}"  ADDIN PROCITE \11\05\19\02\00\00\00\1D{Huang & Ingber 2000 #594210}\00\1D\00(\00\00%Dick 9:Dick's Databases:Dick's Biblio\1BHuang & Ingber 2000 #594210\00\1B\00 {Huang & Ingber 2000 #594210}  QUOTE "{von Dassow, Meir, et al. 2000 #590510}"  ADDIN PROCITE \11\05\19\02\00\00\00'{von Dassow, Meir, et al. 2000 #590510}\00'\00J\00\00%Dick 9:Dick's Databases:Dick's Biblio%von Dassow, Meir, et al. 2000 #590510\00%\00 {von Dassow, Meir, et al. 2000 #590510}. Network models of gene regulation in the nucleus may simply not be capable of explaining embryogenesis. In a recent paper  QUOTE "{Gerner & Sauermann 1999 #176370}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Gerner & Sauermann 1999 #176370}\00!\00u: \00\00%Dick 9:Dick's Databases:Dick's Biblio\1FGerner & Sauermann 1999 #176370\00\1F\00 {Gerner & Sauermann 1999 #176370}, three remarkable features of isolated nuclear matrices of human hematopoietic cells are apparent: Cells of the same kind have nuclear matrices that look the same. Nuclear matrices are the same shape as the nucleus of the intact cell. Cells of different kinds that are closely related by lineage nevertheless can have nuclear matrices of very different shapes. Furthermore, similar nuclear shapes and textures for given blood cell types occur across species  QUOTE "{LeDouarin, Jotereau, et al. 1984 #607750}"  ADDIN PROCITE \11\05\19\02\00\00\00+{Le Douarin, Jotereau, et al. 1984 #607750}\00+\00\00\00%Dick 9:Dick's Databases:Dick's Biblio)Le Douarin, Jotereau, et al. 1984 #607750\00)\00 {LeDouarin, Jotereau, et al. 1984 #607750}  QUOTE "{Ussing & Rosenkilde 1995 #621980}"  ADDIN PROCITE \11\05\19\02\00\00\00"{Ussing & Rosenkilde 1995 #621980}\00"\005\00\00%Dick 9:Dick's Databases:Dick's Biblio Ussing & Rosenkilde 1995 #621980\00 \00 {Ussing & Rosenkilde 1995 #621980}. The practice of pathology is highly dependent on the empirical observation that: The nuclei of most cancer cells, with the exception of some well-differentiated tumors, are abnormally shaped and have an irregular contour, with protrusions, indentations, and grooves  QUOTE "{Ehya 1994 #643220}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Ehya 1994 #643220}\00\13\00\10\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Ehya 1994 #643220\00\11\00 {Ehya 1994 #643220}. Yet little thought has been given to the causal relationship between the overall nuclear structure and the state of the cell. Earlier work seems to have only addressed changes in the size of the nucleus with differentiated state  QUOTE "{D'Amato 1977 #646830}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{D'Amato 1977 #646830}\00\16\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\14D'Amato 1977 #646830\00\14\00 {D'Amato 1977 #646830}. There are many other observations that may have their root cause in an overall architecture to the nucleus: Tugging on one chromosome in a cell allows one to pull out all the chromosomes as clumps on a DNA string  QUOTE "{Maniotis, Bojanowski, et al. 1997 #331600}"  ADDIN PROCITE \11\05\19\02\00\00\00+{Maniotis, Bojanowski, et al. 1997 #331600}\00+\00L: \00\00%Dick 9:Dick's Databases:Dick's Biblio)Maniotis, Bojanowski, et al. 1997 #331600\00)\00 {Maniotis, Bojanowski, et al. 1997 #331600}. This suggests that the genome has an overall one dimensional, perhaps circular topology that folds into a three dimensional structure. Chromosomes have nonrandom positions in the nucleus  QUOTE "{Nagele, Freeman, et al. 1995 #642160}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Nagele, Freeman, et al. 1995 #642160}\00&\00\00\00%Dick 9:Dick's Databases:Dick's Biblio$Nagele, Freeman, et al. 1995 #642160\00$\00 {Nagele, Freeman, et al. 1995 #642160}. When two cells are fused to form a heterokaryon, certain sets of chromosomes are eliminated in the first few cell divisions  QUOTE "{Harris 1995 #212810}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Harris 1995 #212810}\00\15\00l\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Harris 1995 #212810\00\13\00 {Harris 1995 #212810}. Chromosomes in nuclei in epithelia have a specific orientation with respect to the apical and basal surfaces of the cells  QUOTE "{Francis-Lang, Davis, et al. 1996 #810}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Francis-Lang, Davis, et al. 1996 #810}\00'\00\0F\00\00%Dick 9:Dick's Databases:Dick's Biblio%Francis-Lang, Davis, et al. 1996 #810\00%\00 {Francis-Lang, Davis, et al. 1996 #810}. Exposed DNA is typically in only one of two conformations: in reverse transformed cells, it is perinuclear, while in the transformed counterparts, it is in splotches that are seemingly random throughout the nucleus  QUOTE "{Puck & Krystosek 1992 #594660}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Puck & Krystosek 1992 #594660}\00\1F\00I\00\00%Dick 9:Dick's Databases:Dick's Biblio\1DPuck & Krystosek 1992 #594660\00\1D\00 {Puck & Krystosek 1992 #594660}. The severity of an aneuploid phenotype is roughly proportional to the size of the extra chromosome  QUOTE "{Epstein 1986 #642080}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{Epstein 1986 #642080}\00\16\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\14Epstein 1986 #642080\00\14\00 {Epstein 1986 #642080}  QUOTE "{Rasnick & Duesberg 1999 #617340}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Rasnick & Duesberg 1999 #617340}\00!\00{\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FRasnick & Duesberg 1999 #617340\00\1F\00 {Rasnick & Duesberg 1999 #617340}. The consequences of triploidy are generally worse than those of tetraploidy  QUOTE "{Epstein 1986 #642080}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{Epstein 1986 #642080}\00\16\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\14Epstein 1986 #642080\00\14\00 {Epstein 1986 #642080}. These lead to the notion that the nucleus is a discrete state machine  QUOTE "{Nagl 1983 #373360}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Nagl 1983 #373360}\00\13\00'\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Nagl 1983 #373360\00\11\00 {Nagl 1983 #373360}. The Wurfel Model for Nuclear Mechanics A Wurfel (Figure4) is made of 12 cubical blocks of wood connected by a single, topologically circular, taut elastic band. In each block the band comes in through a hole in one face, goes through the center, and exits from an adjacent face. In any of its numerous configurations, the surfaces of a Wurfel fall in one of three categories: a) permanently hidden (faces with elastic); b) exposed; c) sequestered. We take these as roughly corresponding to: a) connections of genes along the chromosomes; b) genes available for transcription; c) silenced genes, respectively  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s \00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660}. In this playful model, transitions between states of nuclear differentiation correspond to configurational changes of a Wurfel. At each transition, new surfaces would be exposed, and some exposed surfaces would be sequestered (cf. selective potentiation of chromatin domains  QUOTE "{Kramer, McCarrey, et al. 1998 #288410}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Kramer, McCarrey, et al. 1998 #288410}\00'\00d"\00\00%Dick 9:Dick's Databases:Dick's Biblio%Kramer, McCarrey, et al. 1998 #288410\00%\00 {Kramer, McCarrey, et al. 1998 #288410}). The energy barrier for going from one configuration to another is roughly the same, so that the "landscape" for a Wurfel consists of multiple peaks of similar height, unlike the energy landscape of a protein, which generally has one major peak, the native state. A problem with Wurfels as a model for the nuclear machine for differentiation is that many states are accessible from a given state, so that it cannot represent our requirement that precisely two new states be accessible from a given state (Figure3). If a given agent, such as a hybrid polar compound  QUOTE "{Richon, Emiliani, et al. 1998 #434620}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Richon, Emiliani, et al. 1998 #434620}\00'\00 \00\00%Dick 9:Dick's Databases:Dick's Biblio%Richon, Emiliani, et al. 1998 #434620\00%\00 {Richon, Emiliani, et al. 1998 #434620}, could swell some parts of the chromatin, while condensing other parts, and another agent would do the reverse, a Wurfel structure would then have precisely two new states to go to. Suppose that the genome transits from differentiation state A to either B or C. Consider state B. If a cell with state B is to differentiate further using the same two agents, then we want B -> D or E. But one of the two agents would just send B back to A. To prevent this, we need to lock up parts of the genome in state B, so that state A is no longer accessible. This is a possible role for methylation, the Rett protein gene MeCP2, which binds to methylated DNA  QUOTE "{Kumar 2000 #621610}"  ADDIN PROCITE \11\05\19\02\00\00\00\14{Kumar 2000 #621610}\00\14\00\11\00\00%Dick 9:Dick's Databases:Dick's Biblio\12Kumar 2000 #621610\00\12\00 {Kumar 2000 #621610}, acetylation, the distinction between heterochromatin and decondensed euchromatin, etc.  QUOTE "{Spencer & Davie 1999 #495150}"  ADDIN PROCITE \11\05\19\02\00\00\00\1E{Spencer & Davie 1999 #495150}\00\1E\00\1F\00\00%Dick 9:Dick's Databases:Dick's Biblio\1CSpencer & Davie 1999 #495150\00\1C\00 {Spencer & Davie 1999 #495150}. With such locking, the agents will alter nuclear state B to D or E, but not back to A. In this way, differentiation could be organized as a binary differentiation tree (Figure 3), rather than as a web  QUOTE "{Kauffman 1973 #269070}"  ADDIN PROCITE \11\05\19\02\00\00\00\17{Kauffman 1973 #269070}\00\17\00%\0C\00\00%Dick 9:Dick's Databases:Dick's Biblio\15Kauffman 1973 #269070\00\15\00 {Kauffman 1973 #269070}. Modelling Chromosomes and the Whole Genome In order to get a better idea of how the Wurfel model might apply to chromosomes, I had sets of alphabet beads redrilled so that the new hole in each bead went at 45o between the centers of two adjacent faces (Figure5a). An elastic string was passed through them and tied under tension, to make the string and beads a tensegrity structure. With the beads in alphabetical order, one can make out the primary sequence despite folding (Figure5b). As with a Wurfel, the alpha-Wurfel has many configurations. However, it has the additional property, probably simply due to the greater length of the chain of beads, of being floppy, i.e., having some unconstrained degrees of rotation. Some of this floppiness can be removed by adding additional elastic rubber bands on each chromosome (Figure 5c). As the whole model is stretched, at first the chromosomes barely respond, due to their lower extensibility (Figure5d). But finally they do change configuration, as the elastic rubber band slips over some of the beads (Figure 5e), a crude model of observed disentanglement  QUOTE "{Poirier, Nemani, et al. 2001 #623350}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Poirier, Nemani, et al. 2001 #623350}\00&\00"\00\00%Dick 9:Dick's Databases:Dick's Biblio$Poirier, Nemani, et al. 2001 #623350\00$\00 {Poirier, Nemani, et al. 2001 #623350}. The alpha-Wurfel model of the genome is a rough working model, but at this point it embodies most of what we know about the mechanics of the whole genome. The mechanics can be studied in detail, by obtaining stress/strain hysteresis curves (cf. individual chromosomes  QUOTE "{Poirier, Eroglu, et al. 2000 #623360}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Poirier, Eroglu, et al. 2000 #623360}\00&\00"\00\00%Dick 9:Dick's Databases:Dick's Biblio$Poirier, Eroglu, et al. 2000 #623360\00$\00 {Poirier, Eroglu, et al. 2000 #623360}), by computed tomography of the genome suspended and rotated between two coaxial micromanipulators, and by inverse methods using forced vibrations  QUOTE "{Gladwell 1986a #629820}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Gladwell 1986a #629820}\00\18\00-\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Gladwell 1986a #629820\00\16\00 {Gladwell 1986a #629820}. These will lead to refined models. If the Wurfel concept is correct, then, when a stretched chromosome changes its configuration, there ought to be a corresponding change in gene expression. Discussion The long history of research on the morphogenetic furrow in the Drosophila eye imaginal disc (starting with  QUOTE "{Poodry, Hall, et al. 1973 #415330}"  ADDIN PROCITE \11\05\19\02\00\00\00#{Poodry, Hall, et al. 1973 #415330}\00#\00\00\00%Dick 9:Dick's Databases:Dick's Biblio!Poodry, Hall, et al. 1973 #415330\00!\00 {Poodry, Hall, et al. 1973 #415330}  QUOTE "{Ready, Hanson, et al. 1976 #429330}"  ADDIN PROCITE \11\05\19\02\00\00\00${Ready, Hanson, et al. 1976 #429330}\00$\00\12\00\00%Dick 9:Dick's Databases:Dick's Biblio"Ready, Hanson, et al. 1976 #429330\00"\00 {Ready, Hanson, et al. 1976 #429330}) is one of neglect to study the wave itself. It has been treated as an epiphenomenon, irrelevant to rather than causal of cell differentiation. In its original formulation  QUOTE "{Wolpert 1969 #572290}"  ADDIN PROCITE \11\05\19\02\00\00\00\16{Wolpert 1969 #572290}\00\16\00A\17\00\00%Dick 9:Dick's Databases:Dick's Biblio\14Wolpert 1969 #572290\00\14\00 {Wolpert 1969 #572290}, positional information theory posited differentiation of all cell types in one grand step: each cell learns its position, uses some sort of lookup table to find out what it is supposed to be as a function of its coordinates, and then changes into that cell type. The contrast with differentiation waves is stark, and testable: a cell has no need to know where it is; it participates in a contraction or expansion wave transmitted from its neighbors; it changes to one of two new cell types, depending on the type of wave. The branches of the differentiation tree representing these alternative cell types correspond to a binary differentiation code (Figure 3), which is the sequence of waves  QUOTE "{Bjrklund & Gordon 1994 #45690}"  ADDIN PROCITE \11\05\19\02\00\00\00 {Bjrklund & Gordon 1994 #45690}\00 \00O<\00\00%Dick 9:Dick's Databases:Dick's Biblio\1EBjrklund & Gordon 1994 #45690\00\1E\00 {Bjrklund & Gordon 1994 #45690}. Problems with gradient theory include: The mechanism of transport of purported morphogens is unknown. Nonlinear effects have to be invoked to obtain sharp boundaries. There is no theory for the spatial arrangement of sources and sinks of morphogens and their timing. We are expected to accept entirely different developmental mechanisms for short and long germ insects  QUOTE "{Akam & Dawes 1992 #5010}"  ADDIN PROCITE \11\05\19\02\00\00\00\19{Akam & Dawes 1992 #5010}\00\19\00q+\00\00%Dick 9:Dick's Databases:Dick's Biblio\17Akam & Dawes 1992 #5010\00\17\00 {Akam & Dawes 1992 #5010}. There is no explanation for why differentiation should be accompanied by contraction or expansion waves. Differentiation wave theory has four major holes: What causes a wave of a given type (expansion or contraction) to be launched at a given place at a given time? What shapes the trajectory of a wave (including where it stops)? What is the signal transduction mechanism from the subcellular apparatus that sustains propagation of a wave through a cell (the cell state splitter, Figure 6:  QUOTE "{Gordon & Brodland 1987a #188950}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Gordon & Brodland 1987a #188950}\00!\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FGordon & Brodland 1987a #188950\00\1F\00 {Gordon & Brodland 1987a #188950}) to the nucleus? Why do some differentiations coincide with thresholds of morphogens  QUOTE "{Gurdon, Standley, et al. 1999 #202350}"  ADDIN PROCITE \11\05\19\02\00\00\00'{Gurdon, Standley, et al. 1999 #202350}\00'\00R \00\00%Dick 9:Dick's Databases:Dick's Biblio%Gurdon, Standley, et al. 1999 #202350\00%\00 {Gurdon, Standley, et al. 1999 #202350}? Possibilities for the binary signal from the mechanochemical cell state splitter to the nucleus include standard signal transduction  QUOTE "{Bjrklund & Gordon 1993b #45670}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Bjrklund & Gordon 1993b #45670}\00!\00\198\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FBjrklund & Gordon 1993b #45670\00\1F\00 {Bjrklund & Gordon 1993b #45670} or a marionette mechanism  QUOTE "{Maniotis, Chen, et al. 1997 #331620}"  ADDIN PROCITE \11\05\19\02\00\00\00%{Maniotis, Chen, et al. 1997 #331620}\00%\00 \00\00%Dick 9:Dick's Databases:Dick's Biblio#Maniotis, Chen, et al. 1997 #331620\00#\00 {Maniotis, Chen, et al. 1997 #331620} involving pushing/pulling of the nucleus. The launching and termination mechanisms of differentiation waves probably correspond to the beginning and end of induction events. Proof that gradients track behind differentiation waves, other than in the Drosophila eye imaginal disc, is not yet forthcoming. The nucleus is a world unto itself, which we are just beginning to explore. Now that we have sequenced the chromosomes, it may be time to complete the sequence of the whole genome, by ascertaining what kind of DNA links the chromosomes  QUOTE "{Maniotis, Chen, et al. 1997 #331620}"  ADDIN PROCITE \11\05\19\02\00\00\00%{Maniotis, Chen, et al. 1997 #331620}\00%\00 \00\00%Dick 9:Dick's Databases:Dick's Biblio#Maniotis, Chen, et al. 1997 #331620\00#\00 {Maniotis, Chen, et al. 1997 #331620}, what the sequence of the chromosomes is along the strand of the whole genome, and if and how the sequence of chromosomes changes during differentiation, in aneuploidy, and in cancer. Wurfels give a working model for the folding of the whole genome, whose configurations may correspond to states of differentiation, and represent one of the codes of biology  QUOTE "{Barbieri 2001a #615270}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Barbieri 2001a #615270}\00\18\00e"\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Barbieri 2001a #615270\00\16\00 {Barbieri 2001a #615270}. This is a very different approach from that of basins of attraction in networks of regulatory genes. The full truth may lie somewhere in between. The derivation of specific cell types from stem cells has aroused great interest recently. But the techniques for causing one cell type to turn into another are a hodge podge, and there is little or no fundamental understanding or control of what is going on  QUOTE "{Shamblott, Edwards, et al. 2000 #647350}"  ADDIN PROCITE \11\05\19\02\00\00\00){Shamblott, Edwards, et al. 2000 #647350}\00)\009 \00\00%Dick 9:Dick's Databases:Dick's Biblio'Shamblott, Edwards, et al. 2000 #647350\00'\00 {Shamblott, Edwards, et al. 2000 #647350}. If a Wurfel-like state of the genome exists for each cell type, then we might be able to rationalize cell type transitions. We postulated that germ cells in urodele amphibians are cells missed by early differentiation waves  QUOTE "{Bjrklund & Gordon 1994 #45690}"  ADDIN PROCITE \11\05\19\02\00\00\00 {Bjrklund & Gordon 1994 #45690}\00 \00O<\00\00%Dick 9:Dick's Databases:Dick's Biblio\1EBjrklund & Gordon 1994 #45690\00\1E\00 {Bjrklund & Gordon 1994 #45690}. Perhaps, at any stage of differentiation, stem cells are precisely those few cells at the margins of embryonic tissues that are missed by both the expansion and contraction wave that other cells in their tissue participated in. They are stuck, so to speak, with incomplete differentiation codes (Figure 3). If stem cells (including embryonic stem cells), were to be taken mechanically or biochemically through a particular sequence of contractions and expansions corresponding to the subsequent differentiation code (Figure3), then, if differentiation wave theory works, it should be possible to create any subsequent tissue from them (predifferentiation:  QUOTE "{Keirstead 2001 #648530}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Keirstead 2001 #648530}\00\18\00F\00\01\00%Dick 9:Dick's Databases:Dick's Biblio\16Keirstead 2001 #648530\00\16\00 {Keirstead 2001 #648530}). This could be automated as a  tissue synthesizer for any needed replacement tissue, for any individual person. Summary Two contesting paradigms for development The prediction  QUOTE "{Gordon & Brodland 1987a #188950}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Gordon & Brodland 1987a #188950}\00!\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FGordon & Brodland 1987a #188950\00\1F\00 {Gordon & Brodland 1987a #188950} and discovery  QUOTE "{Brodland, Gordon, et al. 1994 #62320}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Brodland, Gordon, et al. 1994 #62320}\00&\00!\00\00%Dick 9:Dick's Databases:Dick's Biblio$Brodland, Gordon, et al. 1994 #62320\00$\00 {Brodland, Gordon, et al. 1994 #62320}  QUOTE "{Gordon, Bjrklund, et al. 1994 #188900}"  ADDIN PROCITE \11\05\19\02\00\00\00({Gordon, Bjrklund, et al. 1994 #188900}\00(\00V%\00\00%Dick 9:Dick's Databases:Dick's Biblio&Gordon, Bjrklund, et al. 1994 #188900\00&\00 {Gordon, Bjrklund, et al. 1994 #188900} of differentiation waves  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660} has driven me to conclusions contrary to the prevailing paradigm that gradients of positional information are key to embryonic development: Proposition 35: Positional information does not exist. Proposition 242: Gradients of morphogens are irrelevant to differentiation  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s \00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660}. This might seem a brash position to take, given what is regarded by some as overwhelming evidence for positional information. But the evidence against positional information via gradients is also persuasive, especially in the existence of polyembryonic wasps  QUOTE "{Grbic, Nagy, et al. 1998 #194360}"  ADDIN PROCITE \11\05\19\02\00\00\00"{Grbic, Nagy, et al. 1998 #194360}\00"\00" H"\00\00%Dick 9:Dick's Databases:Dick's Biblio Grbic, Nagy, et al. 1998 #194360\00 \00 {Grbic, Nagy, et al. 1998 #194360} and the ability of chick embryo fragments to form whole chick embryos  QUOTE "{Perrimon & Stern 1999 #406630}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Perrimon & Stern 1999 #406630}\00\1F\00b\1A\00\00%Dick 9:Dick's Databases:Dick's Biblio\1DPerrimon & Stern 1999 #406630\00\1D\00 {Perrimon & Stern 1999 #406630}. Differentiation waves provide a clear, testable, alternative hypothesis. I see them as the first event in each step of differentiation, rather than regarding the erection of a gradient as the first event. Two contesting paradigms for the genome in development Ever since Morgan  QUOTE "{Morgan 1934 #365150}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Morgan 1934 #365150}\00\15\00Q\10\00\00%Dick 9:Dick's Databases:Dick's Biblio\13Morgan 1934 #365150\00\13\00 {Morgan 1934 #365150} recognized that differentiated cells in an organism could have different  batteries of genes active, we have been without a mechanism for how this comes about and is maintained. Today two paradigms are being considered: 1) differentiation is a matter of networks of gene interactions settling down into one of the available stable network states (basins of attraction)  QUOTE "{Delbrck 1949a #118650}"  ADDIN PROCITE \11\05\19\02\00\00\00\18{Delbrck 1949a #118650}\00\18\00S\00\00%Dick 9:Dick's Databases:Dick's Biblio\16Delbrck 1949a #118650\00\16\00 {Delbrck 1949a #118650}  QUOTE "{Kauffman 1993 #269200}"  ADDIN PROCITE \11\05\19\02\00\00\00\17{Kauffman 1993 #269200}\00\17\00v5\00\00%Dick 9:Dick's Databases:Dick's Biblio\15Kauffman 1993 #269200\00\15\00 {Kauffman 1993 #269200}; 2) the genome or nucleus is a machine that has multiple physical states (conformations) which expose and sequester different subsets of genes  QUOTE "{Gordon 1999 #188660}"  ADDIN PROCITE \11\05\19\02\00\00\00\15{Gordon 1999 #188660}\00\15\00s \00\00%Dick 9:Dick's Databases:Dick's Biblio\13Gordon 1999 #188660\00\13\00 {Gordon 1999 #188660}. While I have shown how the differentiation wave paradigm can be linked to the nucleus as machine paradigm in the working model in Figure 6, other pairs of paradigms for cellular and nuclear differentiation during embryogenesis may be plausible (Table 1). The postulated sequence of events, for the Wave & Wurfel pair of paradigms for differentiation that I favor, is thus: A cell receives a signal to expand or contract from a neighboring cell that has just done the same. This is the differentiation wave. Its cell state splitter expands or contracts (sending the wave on to the next cell in the tissue). Signal transduction from the cell state splitter to the nucleus sends a one bit signal to the nucleus indicating which kind of wave the cell just participated in. The genome machine changes to one of two new discrete configurations via a Wurfel-like mechanism. Some genes are exposed (made available for transcription) and others are sequestered as a result of the change in genome configuration. The change in gene products changes the phenotype of the cell. Processes such as methylation prevent the new genome configuration from flipping back to the old one in the subsequent differentiation. The cell state splitter is readied for the next wave. The experimental challenge in choosing between the four pairs of paradigms (Table 1) is to combine the fine methods of molecular developmental biology with time lapse, mechanics of cells and genomes, and analysis of nuclear structure and function, at the critical moments when cells in embryos change from one differentiated state to another. Acknowledgements I would like to thank Andrew J. Maniotis for pointing out that the Wurfel model for the nucleus in differentiation makes the nucleus a machine for differentiation, and Ian Paul and Sean Billing for assistance in constructing the Wurfel-like models. Critical readings were kindly done by Andrew J. Maniotis, Alexei G. Desnitski, John Parkinson, Christopher Gerner, Adam S. Wilkins and Natalie K. Bjrklund. d  Figure1. Models for movement of the morphogen molecule. a) Diffusion through the extracellular space. b) Planar transcytosis. c) Displacement during growth. (From  QUOTE "{Tabata 2001 #1570}"  ADDIN PROCITE \11\05\19\02\00\00\00\13{Tabata 2001 #1570}\00\13\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\11Tabata 2001 #1570\00\11\00 {Tabata 2001 #1570} with the authors permission.) d) A differentiation wave (contraction, as illustrated, or expansion wave) initiates synthesis of a purported morphogen, which accumulates over time in each cell. This produces a gradient from cell to cell in the wake of the wave. Note that for differentiation waves there is no requirement for intercellular transport of the morphogen, and that morphogen synthesis is a consequence of differentiation (or, at least, determination), not its cause.  Figure2. The ectoderm contraction wave appears at early gastrulation at a focus above the dorsal lip of the blastopore, which was classically regarded as the organizer  QUOTE "{Spemann & Mangold 1924c #631080}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Spemann & Mangold 1924c #631080}\00!\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FSpemann & Mangold 1924c #631080\00\1F\00 {Spemann & Mangold 1924c #631080}. The initiation focus is probably underlain by pharyngeal endoderm  QUOTE "{Gordon & Brodland 1987a #188950}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Gordon & Brodland 1987a #188950}\00!\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FGordon & Brodland 1987a #188950\00\1F\00 {Gordon & Brodland 1987a #188950}. This composite is reconstructed from a number of partial time lapse sequences of the wave  QUOTE "{Brodland, Gordon, et al. 1994 #62320}"  ADDIN PROCITE \11\05\19\02\00\00\00&{Brodland, Gordon, et al. 1994 #62320}\00&\00!\00\00%Dick 9:Dick's Databases:Dick's Biblio$Brodland, Gordon, et al. 1994 #62320\00$\00 {Brodland, Gordon, et al. 1994 #62320}. Times and dimensions are given for the axolotl, Ambystoma mexicanum, at 20oC. The curved lines show the bottom (trough) of the furrow at 1 hr intervals. Its manner of vanishing, at the end of gastrulation, suggests that it is a solitary wave in an active medium. From  QUOTE "{Gordon, Bjrklund, et al. 1994 #188900}"  ADDIN PROCITE \11\05\19\02\00\00\00({Gordon, Bjrklund, et al. 1994 #188900}\00(\00V%\00\00%Dick 9:Dick's Databases:Dick's Biblio&Gordon, Bjrklund, et al. 1994 #188900\00&\00 {Gordon, Bjrklund, et al. 1994 #188900} with permission.  Figure3. Each arrow of a differentiation tree corresponds to an embryonic or terminal tissue, named here at its arrowhead. Each tissue is generally split into two tissues by a pair of spatially complementary waves, one contraction wave and one expansion wave. The differentiation tree for an organism is thus its tissue lineage tree. This is the differentiation tree of Ambystoma mexicanum, from a single fertilized egg through the end of gastrulation. The differentiation code for each tissue is shown, i.e., the sequence of observed expansion (E) and contraction (C) waves that the cell has so far participated in. For an accurate depiction of developmental time, which proceeds top down in this figure, see  QUOTE "{Gordon, Bjrklund, et al. 1994 #188900}"  ADDIN PROCITE \11\05\19\02\00\00\00({Gordon, Bjrklund, et al. 1994 #188900}\00(\00V%\00\00%Dick 9:Dick's Databases:Dick's Biblio&Gordon, Bjrklund, et al. 1994 #188900\00&\00 {Gordon, Bjrklund, et al. 1994 #188900}. From  QUOTE "{Bjrklund & Gordon 1994 #45690}"  ADDIN PROCITE \11\05\19\02\00\00\00 {Bjrklund & Gordon 1994 #45690}\00 \00O<\00\00%Dick 9:Dick's Databases:Dick's Biblio\1EBjrklund & Gordon 1994 #45690\00\1E\00 {Bjrklund & Gordon 1994 #45690} with permission.   Figure4. A Wurfel in three configurations of many (also called Rainbow Twist Cubes: Pappa Geppetto's Toys Victoria Ltd., Victoria, Canada, with permission of Manhattan Toy). a  b  c  d  e  Figure5. a) An extended alpha-Wurfel arrangement of alphabet beads, showing that the string enters and exits at adjacent faces on each block. b) A clumpy configuration, in which specific faces of the alpha-Wurfel blocks are sequestered or exposed, representing gene silencing and activation, respectively. The clump could represent one chromosome along the genome. Note that the beads used for this model have no adhesions between them, other than the elastic that goes through them. To maintain a clump, we have to assume that touching faces have some sort of bond between them. This was not necessary for the short loop of 12 blocks in a Wurfel (Figure4), but in an open chain, the single elastic string is insufficient to hold a given configuration by itself. c) An elastic band is tied around each Wurfel-like chromosome in a genome, to bind it together and give it some elasticity. The linker DNA is represented by highly elastic coils (John Marko & Andrew Maniotis, personal communication), which are in their relaxed state here. d) When the genome is stretched, the chromosomes hardly respond. e) The genome is overstretched, resulting in slippage of the elastic rubber band clumping a chromosome.  Figure6. The cell state splitter is a nonlinear tensegrity apparatus (capable of snapping between configurations) at the apical end of a cell. It consists of a contractile microfilament ring (MF) in radial tension with a subtending annular mat of apical microtubules (MT). An intermediate filament ring (IF) prevents immediate contraction or expansion of the cell state splitter by keeping it in a mechanically metastable state  QUOTE "{Gordon & Brodland 1987a #188950}"  ADDIN PROCITE \11\05\19\02\00\00\00!{Gordon & Brodland 1987a #188950}\00!\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1FGordon & Brodland 1987a #188950\00\1F\00 {Gordon & Brodland 1987a #188950}  QUOTE "{Martin & Gordon 1997b #336690}"  ADDIN PROCITE \11\05\19\02\00\00\00\1F{Martin & Gordon 1997b #336690}\00\1F\00\00\00%Dick 9:Dick's Databases:Dick's Biblio\1DMartin & Gordon 1997b #336690\00\1D\00 {Martin & Gordon 1997b #336690}. Cell A participates in either a contraction wave (MF based), which turns it into a cell of type B, or an expansion wave (MT based), which turns it into a cell of type C. The thick arrows represent one branch of the organisms differentiation tree (Figure 3). The wave is transmitted to the next cell(s) in the epithelium (Figure 1d). Signal transduction sends a single bit of information from the cell state splitter to the nucleus which indicates whether the cell just participated in a contraction wave or an expansion wave. The Wurfel-like structure of the genome changes to one of two new configurations. This results in a change in gene expression, represented by the newly exposed and sequestered surfaces of the Wurfel. The change in genome structure may be reflected by a change in shape of the nucleus. Reference List 1. Akam ME, Dawes R. Insect segmentation: more than one way to slice an egg. Curr. Biol. 1992; 2:395-398. 2. Almirantis Y, Papageorgiou S. Modes of morphogen cooperation for limb formation in vertebrates and insects. J Theor Biol 1999; 199:235-242. 3. Barbieri, M. (2001a). The Organic Codes: The Birth of Semantic Biology, Ancona, Italy: Casa editrice Pequod. 4. Bell, E. (2000). Tissue engineering in perspective. In: Lanza, R.P., R. Langer & J. 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ThreeTabB ThreeTabCTwoTabCTwoTabDMPEG Layer-3 Decompressor MPEG Layer-3(MPEG Layer-3 Decompressor (m/8kbit/8kHz)-MPEG Layer-3 Decompressor (m/8kbit/11.025kHz))MPEG Layer-3 Decompressor (m/8kbit/12kHz))MPEG Layer-3 Decompressor (m/8kbit/16kHz),MPEG Layer-3 Decompressor (m/8kbit/22.05kHz))MPEG Layer-3 Decompressor (m/8kbit/24kHz))MPEG Layer-3 Decompressor (m/8kbit/32kHz)+MPEG Layer-3 Decompressor (m/8kbit/44.1kHz))MPEG Layer-3 Decompressor (m/8kbit/48kHz))MPEG Layer-3 Decompressor (m/16kbit/8kHz).MPEG Layer-3 Decompressor (m/16kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/16kbit/12kHz)*MPEG Layer-3 Decompressor (m/16kbit/16kHz)-MPEG Layer-3 Decompressor (m/16kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/16kbit/24kHz)*MPEG Layer-3 Decompressor (m/16kbit/32kHz),MPEG Layer-3 Decompressor (m/16kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/16kbit/48kHz))MPEG Layer-3 Decompressor (m/18kbit/8kHz).MPEG Layer-3 Decompressor (m/18kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/18kbit/12kHz)*MPEG Layer-3 Decompressor (m/18kbit/16kHz)-MPEG Layer-3 Decompressor (m/18kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/18kbit/24kHz)*MPEG Layer-3 Decompressor (m/18kbit/32kHz),MPEG Layer-3 Decompressor (m/18kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/18kbit/48kHz))MPEG Layer-3 Decompressor (m/20kbit/8kHz).MPEG Layer-3 Decompressor (m/20kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/20kbit/12kHz)*MPEG Layer-3 Decompressor (m/20kbit/16kHz)-MPEG Layer-3 Decompressor (m/20kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/20kbit/24kHz)*MPEG Layer-3 Decompressor (m/20kbit/32kHz),MPEG Layer-3 Decompressor (m/20kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/20kbit/48kHz))MPEG Layer-3 Decompressor (m/24kbit/8kHz).MPEG Layer-3 Decompressor (m/24kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/24kbit/12kHz)*MPEG Layer-3 Decompressor (m/24kbit/16kHz)-MPEG Layer-3 Decompressor (m/24kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/24kbit/24kHz)*MPEG Layer-3 Decompressor (m/24kbit/32kHz),MPEG Layer-3 Decompressor (m/24kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/24kbit/48kHz))MPEG Layer-3 Decompressor (m/32kbit/8kHz).MPEG Layer-3 Decompressor (m/32kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/32kbit/12kHz)*MPEG Layer-3 Decompressor (m/32kbit/16kHz)-MPEG Layer-3 Decompressor (m/32kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/32kbit/24kHz)*MPEG Layer-3 Decompressor (m/32kbit/32kHz),MPEG Layer-3 Decompressor (m/32kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/32kbit/48kHz))MPEG Layer-3 Decompressor (m/48kbit/8kHz).MPEG Layer-3 Decompressor (m/48kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/48kbit/12kHz)*MPEG Layer-3 Decompressor (m/48kbit/16kHz)-MPEG Layer-3 Decompressor (m/48kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/48kbit/24kHz)*MPEG Layer-3 Decompressor (m/48kbit/32kHz),MPEG Layer-3 Decompressor (m/48kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/48kbit/48kHz))MPEG Layer-3 Decompressor (m/56kbit/8kHz).MPEG Layer-3 Decompressor (m/56kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/56kbit/12kHz)*MPEG Layer-3 Decompressor (m/56kbit/16kHz)-MPEG Layer-3 Decompressor (m/56kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/56kbit/24kHz)*MPEG Layer-3 Decompressor (m/56kbit/32kHz),MPEG Layer-3 Decompressor (m/56kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/56kbit/48kHz))MPEG Layer-3 Decompressor (m/64kbit/8kHz).MPEG Layer-3 Decompressor (m/64kbit/11.025kHz)*MPEG Layer-3 Decompressor (m/64kbit/12kHz)*MPEG Layer-3 Decompressor (m/64kbit/16kHz)-MPEG Layer-3 Decompressor (m/64kbit/22.05kHz)*MPEG Layer-3 Decompressor (m/64kbit/24kHz)*MPEG Layer-3 Decompressor (m/64kbit/32kHz),MPEG Layer-3 Decompressor (m/64kbit/44.1kHz)*MPEG Layer-3 Decompressor (m/64kbit/48kHz)(MPEG Layer 3 Decompressor (s/8kbit/8kHz)-MPEG Layer 3 Decompressor (s/8kbit/11.025kHz))MPEG Layer 3 Decompressor (s/8kbit/12kHz))MPEG Layer 3 Decompressor (s/8kbit/16kHz),MPEG Layer 3 Decompressor (s/8kbit/22.05kHz))MPEG Layer 3 Decompressor (s/8kbit/24kHz))MPEG Layer 3 Decompressor (s/8kbit/32kHz)+MPEG Layer 3 Decompressor (s/8kbit/44.1kHz))MPEG Layer 3 Decompressor (s/8kbit/48kHz))MPEG Layer 3 Decompressor (s/16kbit/8kHz).MPEG Layer 3 Decompressor (s/16kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/16kbit/12kHz)*MPEG Layer 3 Decompressor (s/16kbit/16kHz)-MPEG Layer 3 Decompressor (s/16kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/16kbit/24kHz)*MPEG Layer 3 Decompressor (s/16kbit/32kHz),MPEG Layer 3 Decompressor (s/16kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/16kbit/48kHz))MPEG Layer 3 Decompressor (s/18kbit/8kHz).MPEG Layer 3 Decompressor (s/18kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/18kbit/12kHz)*MPEG Layer 3 Decompressor (s/18kbit/16kHz)-MPEG Layer 3 Decompressor (s/18kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/18kbit/24kHz)*MPEG Layer 3 Decompressor (s/18kbit/32kHz),MPEG Layer 3 Decompressor (s/18kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/18kbit/48kHz))MPEG Layer 3 Decompressor (s/20kbit/8kHz).MPEG Layer 3 Decompressor (s/20kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/20kbit/12kHz)*MPEG Layer 3 Decompressor (s/20kbit/16kHz)-MPEG Layer 3 Decompressor (s/20kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/20kbit/24kHz)*MPEG Layer 3 Decompressor (s/20kbit/32kHz),MPEG Layer 3 Decompressor (s/20kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/20kbit/48kHz))MPEG Layer 3 Decompressor (s/24kbit/8kHz).MPEG Layer 3 Decompressor (s/24kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/24kbit/12kHz)*MPEG Layer 3 Decompressor (s/24kbit/16kHz)-MPEG Layer 3 Decompressor (s/24kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/24kbit/24kHz)*MPEG Layer 3 Decompressor (s/24kbit/32kHz),MPEG Layer 3 Decompressor (s/24kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/24kbit/48kHz))MPEG Layer 3 Decompressor (s/32kbit/8kHz).MPEG Layer 3 Decompressor (s/32kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/32kbit/12kHz)*MPEG Layer 3 Decompressor (s/32kbit/16kHz)-MPEG Layer 3 Decompressor (s/32kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/32kbit/24kHz)*MPEG Layer 3 Decompressor (s/32kbit/32kHz),MPEG Layer 3 Decompressor (s/32kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/32kbit/48kHz))MPEG Layer 3 Decompressor (s/48kbit/8kHz).MPEG Layer 3 Decompressor (s/48kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/48kbit/12kHz)*MPEG Layer 3 Decompressor (s/48kbit/16kHz)-MPEG Layer 3 Decompressor (s/48kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/48kbit/24kHz)*MPEG Layer 3 Decompressor (s/48kbit/32kHz),MPEG Layer 3 Decompressor (s/48kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/48kbit/48kHz))MPEG Layer 3 Decompressor (s/56kbit/8kHz).MPEG Layer 3 Decompressor (s/56kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/56kbit/12kHz)*MPEG Layer 3 Decompressor (s/56kbit/16kHz)-MPEG Layer 3 Decompressor (s/56kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/56kbit/24kHz)*MPEG Layer 3 Decompressor (s/56kbit/32kHz),MPEG Layer 3 Decompressor (s/56kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/56kbit/48kHz))MPEG Layer 3 Decompressor (s/64kbit/8kHz).MPEG Layer 3 Decompressor (s/64kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/64kbit/12kHz)*MPEG Layer 3 Decompressor (s/64kbit/16kHz)-MPEG Layer 3 Decompressor (s/64kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/64kbit/24kHz)*MPEG Layer 3 Decompressor (s/64kbit/32kHz),MPEG Layer 3 Decompressor (s/64kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/64kbit/48kHz))MPEG Layer 3 Decompressor (s/96kbit/8kHz).MPEG Layer 3 Decompressor (s/96kbit/11.025kHz)*MPEG Layer 3 Decompressor (s/96kbit/12kHz)*MPEG Layer 3 Decompressor (s/96kbit/16kHz)-MPEG Layer 3 Decompressor (s/96kbit/22.05kHz)*MPEG Layer 3 Decompressor (s/96kbit/24kHz)*MPEG Layer 3 Decompressor (s/96kbit/32kHz),MPEG Layer 3 Decompressor (s/96kbit/44.1kHz)*MPEG Layer 3 Decompressor (s/96kbit/48kHz)*MPEG Layer 3 Decompressor (s/112kbit/8kHz)/MPEG Layer 3 Decompressor (s/112kbit/11.025kHz)+MPEG Layer 3 Decompressor (s/112kbit/12kHz)+MPEG Layer 3 Decompressor (s/112kbit/16kHz).MPEG Layer 3 Decompressor (s/112kbit/22.05kHz)+MPEG Layer 3 Decompressor (s/112kbit/24kHz)+MPEG Layer 3 Decompressor (s/112kbit/32kHz)-MPEG Layer 3 Decompressor (s/112kbit/44.1kHz)+MPEG Layer 3 Decompressor 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