205:
2742:
353:). Mesenchymal cells typically leave the epithelial tissue as a consequence of changes in cell adhesive and contractile properties. Following epithelial-mesenchymal transition, cells can migrate away from an epithelium and then associate with other similar cells in a new location. In plants, cellular morphogenesis is tightly linked to the chemical composition and the mechanical properties of the cell wall.
184:, Turing correctly predicted a mechanism of morphogenesis, the diffusion of two different chemical signals, one activating and one deactivating growth, to set up patterns of development, decades before the formation of such patterns was observed. The fuller understanding of the mechanisms involved in actual organisms required the discovery of the structure of
365:. For instance, homotypic cell adhesion can maintain boundaries between groups of cells that have different adhesion molecules. Furthermore, cells can sort based upon differences in adhesion between the cells, so even two populations of cells with different levels of the same adhesion molecule can sort out. In
309:
369:
cells that have the strongest adhesion move to the center of a mixed aggregates of cells. Moreover, cell-cell adhesion is often modulated by cell contractility, which can exert forces on the cell-cell contacts so that two cell populations with equal levels of the same adhesion molecule can sort out.
572:
interact with each other in a characteristic sequence. Maintaining an appropriate balance in the amounts of each of these proteins produced during viral infection appears to be critical for normal phage T4 morphogenesis. Phage T4 encoded proteins that determine virion structure include major
332:
At a tissue level, ignoring the means of control, morphogenesis arises because of cellular proliferation and motility. Morphogenesis also involves changes in the cellular structure or how cells interact in tissues. These changes can result in tissue elongation, thinning, folding, invasion or
573:
structural components, minor structural components and non-structural proteins that catalyze specific steps in the morphogenesis sequence. Phage T4 morphogenesis is divided into three independent pathways: the head, the tail and the long tail fibres as detailed by Yap and
Rossman.
465:. There are often periodic pulses of contraction in embryonic morphogenesis. A model called the cell state splitter involves alternating cell contraction and expansion, initiated by a bistable organelle at the apical end of each cell. The organelle consists of
287:
switch off their cell-to-cell adhesion, become migratory, and take up new positions within an embryo where they again activate specific cell adhesion proteins and form new tissues and organs. Developmental signaling pathways implicated in morphogenesis include
121:
Morphogenesis is a mechanical process involving forces that generate mechanical stress, strain, and movement of cells, and can be induced by genetic programs according to the spatial patterning of cells within tissues. Abnormal morphogenesis is called
1928:
Snustad, D.Peter (1968). "Dominance interactions in
Escherichia coli cells mixedly infected with bacteriophage T4D wild-type and amber mutants and their possible implications as to type of gene-product function: Catalytic vs. Stoichiometric".
337:. Cell "sorting out" consists of cells moving so as to sort into clusters that maximize contact between cells of the same type. The ability of cells to do this has been proposed to arise from differential cell adhesion by
2150:
Sudhakaran, Shyam; Grbic, Djordje; Li, Siyan; Katona, Adam; Najarro, Elias; Glanois, Claire; Risi, Sebastian (2021). "Growing 3D Artefacts and
Functional Machines with Neural Cellular Automata".
730:
Thompson's book is often cited. An abridged version, comprising 349 pages, remains in print and readily obtainable. An unabridged version, comprising 1116 pages, has also been published.
611:
in the twenty-first century enabled the simulation of relatively complex morphogenesis models. In 2020, such a model was proposed where cell growth and differentiation is that of a
247:
are soluble molecules that can diffuse and carry signals that control cell differentiation via concentration gradients. Morphogens typically act through binding to specific protein
515:. The branching is a result of the tip of each bronchiolar tube bifurcating, and the process of branching morphogenesis forms the bronchi, bronchioles, and ultimately the alveoli.
370:
The molecules responsible for adhesion are called cell adhesion molecules (CAMs). Several types of cell adhesion molecules are known and one major class of these molecules are
442:
can contract different parts of the cytoplasm to change its shape or structure. Myosin-driven contractility in embryonic tissue morphogenesis is seen during the separation of
477:
of contraction or expansion over presumptive tissues that determine cell type and is followed by cell differentiation. The cell state splitter was first proposed to explain
361:
During embryonic development, cells are restricted to different layers due to differential affinities. One of the ways this can occur is when cells share the same cell-to-
378:(found on many epithelial cells) binds preferentially to other E-cadherin molecules. Mesenchymal cells usually express other cadherin types such as N-cadherin.
817:
1843:
626:
A similar model to the one described above was subsequently extended to generate three-dimensional structures, and was demonstrated in the video game
2015:
957:
176:(1952). Where Thompson explained animal body shapes as being created by varying rates of growth in different directions, for instance to create the
374:. There are dozens of different cadherins that are expressed on different cell types. Cadherins bind to other cadherins in a like-to-like manner:
2772:
473:
in mechanical opposition. It responds to local mechanical perturbations caused by morphogenetic movements. These then trigger traveling
1851:
144:
Some of the earliest ideas and mathematical descriptions on how physical processes and constraints affect biological growth, and hence
406:
are used to bind to the ECM. Integrins bind extracellularly to fibronectin, laminin, or other ECM components, and intracellularly to
402:
are major ECM molecules that are secreted and assembled into sheets, fibers, and gels. Multisubunit transmembrane receptors called
267:
of other genes; in turn, these secondary gene products can regulate the expression of still other genes in a regulatory cascade of
2721:
390:(ECM) is involved in keeping tissues separated, providing structural support or providing a structure for cells to migrate on.
135:
1209:
1823:
1647:
1622:
1225:
936:
905:
350:
2686:
172:
110:
is an example of highly abnormal and pathological tissue morphogenesis. Morphogenesis also describes the development of
2531:
2259:
230:
208:
2231:
2200:
Artificial Life model of multicellular morphogenesis with autonomously generated gradients for positional information
1680:
866:
2488:
2274:
2269:
342:
438:
Tissues can change their shape and separate into distinct layers via cell contractility. Just as in muscle cells,
1876:
705:
474:
1369:
Angst, B.; Marcozzi, C.; Magee, A. (February 2001). "The cadherin superfamily: diversity in form and function".
1151:
Duran-Nebreda, Salva; Pla, Jordi; Vidiella, Blai; Piñero, Jordi; Conde-Pueyo, Nuria; Solé, Ricard (2021-01-15).
2762:
31:
2665:
2767:
2516:
662:
17:
1706:"Regulation of mammary gland branching morphogenesis by the extracellular matrix and its remodeling enzymes"
1406:"Regulation of mammary gland branching morphogenesis by the extracellular matrix and its remodeling enzymes"
157:
710:
2483:
687:
632:, whose block-based nature made it particularly expedient for the simulation of 3D cellular automatons.
114:
life forms that do not have an embryonic stage in their life cycle. Morphogenesis is essential for the
2732:
2675:
2541:
2373:
1242:
1094:"Modeling mammary organogenesis from biological first principles: Cells and their physical constraints"
1045:"Candidate regulators of mammary branching morphogenesis identified by genome-wide transcript analysis"
897:
799:
642:
293:
2368:
647:
752:
891:
793:
692:
346:
264:
80:
2609:
2546:
2521:
2404:
2324:
1511:
Gordon, Richard; Brodland, G. Wayne (1987). "The cytoskeletal mechanics of brain morphogenesis".
700:
268:
248:
238:
1462:"Key stages in mammary gland development: the cues that regulate ductal branching morphogenesis"
615:
with parametrized rules. As the rules' parameters are differentiable, they can be trained with
2594:
2493:
2478:
2447:
2348:
2060:"Positional information and reaction-diffusion: two big ideas in developmental biology combine"
451:
426:
cascades when binding to the ECM. A well-studied example of morphogenesis that involves ECM is
362:
103:
271:. At the end of this cascade are classes of molecules that control cellular behaviors such as
2705:
2599:
2526:
2425:
2363:
2358:
2289:
2224:
1813:
1334:
Hulpiau, P.; van Roy, F. (February 2009). "Molecular evolution of the cadherin superfamily".
1100:. From the Century of the Genome to the Century of the Organism: New Theoretical Approaches.
652:
545:
can result from disruption of normal morphogenesis, including both tumor formation and tumor
289:
212:
87:
68:
2614:
2457:
2452:
2334:
2329:
2316:
2024:
966:
834:
608:
562:
527:
387:
252:
234:
8:
2710:
2204:
753:"Mechanical stress initiates and sustains the morphogenesis of wavy leaf epidermal cells"
751:
Bidhendi, Amir J.; Altartouri, Bara; Gosselin, Frédérick P.; Geitmann, Anja (July 2019).
423:
321:
2028:
1811:
970:
838:
2399:
2151:
2132:
2040:
1990:
1965:
1868:
1789:
1756:
1582:
1555:
1536:
1488:
1461:
1311:
1286:
1185:
1152:
1128:
1105:
1093:
1069:
1044:
1020:
995:
858:
612:
502:
162:
52:
1732:
1705:
1672:
1432:
1405:
553:
dysfunction can result in increased cancer risk due to disturbed morphogen signaling.
2660:
2440:
2435:
2344:
2136:
2091:
1995:
1946:
1942:
1910:
1906:
1860:
1819:
1794:
1776:
1737:
1686:
1676:
1643:
1618:
1587:
1528:
1493:
1437:
1386:
1351:
1316:
1267:
1221:
1214:
1190:
1172:
1133:
1119:
1074:
1025:
932:
901:
850:
825:
774:
657:
623:. This model was limited to the generation of pictures, and is thus bi-dimensional.
601:
338:
189:
76:
2644:
2109:
Mordvintsev, Alexander; Randazzo, Ettore; Niklasson, Eyvind; Levin, Michael (2020).
1243:"Relating the mechanical properties of the primary plant cell wall to morphogenesis"
996:"Orientation of Turing-like Patterns by Morphogen Gradients and Tissue Anisotropies"
862:
333:
separation of one tissue into distinct layers. The latter case is often referred as
2670:
2639:
2634:
2589:
2217:
2122:
2081:
2071:
2032:
1985:
1977:
1938:
1902:
1872:
1784:
1768:
1727:
1717:
1668:
1610:
1577:
1567:
1540:
1520:
1483:
1473:
1427:
1417:
1378:
1343:
1306:
1298:
1257:
1180:
1164:
1123:
1115:
1064:
1056:
1015:
1007:
974:
842:
764:
620:
616:
582:
512:
145:
123:
60:
2044:
2680:
2584:
2394:
1347:
769:
682:
415:
411:
177:
2059:
1812:
Jay R. Harris; Marc E. Lippman; C. Kent
Osborne; Monica Morrow (28 March 2012).
1772:
619:, a technique which has been highly optimized in recent years due to its use in
94:. Morphogenesis can take place also in a mature organism, such as in the normal
2746:
2209:
1011:
813:
594:
593:'s 1952 paper, "The chemical basis of morphogenesis", a model now known as the
447:
272:
139:
56:
41:
1893:
Floor, Erik (1970). "Interaction of morphogenetic genes of bacteriophage T4".
1572:
1168:
534:
during puberty and is further refined in line with mammary gland development.
328:
than the green cells. The mixed culture forms large multi-cellular aggregates.
2756:
2648:
2562:
2264:
1780:
1176:
1153:"Synthetic Lateral Inhibition in Periodic Pattern Forming Microbial Colonies"
677:
550:
530:, but the branching formation of the duct system begins later in response to
523:
470:
427:
407:
276:
2177:
Morphogenesis: The
Cellular and Molecular Processes of Developmental Anatomy
846:
818:"Morphogenetic Cell Movements: Diversity from Modular Mechanical Properties"
86:
The process controls the organized spatial distribution of cells during the
2473:
2294:
2284:
2127:
2110:
2095:
2036:
1999:
1864:
1798:
1741:
1690:
1667:. Current Topics in Developmental Biology. Vol. 81. pp. 291–310.
1591:
1497:
1441:
1390:
1355:
1320:
1271:
1194:
1137:
1092:
Montévil, Maël; Speroni, Lucia; Sonnenschein, Carlos; Soto, Ana M. (2016).
1078:
1029:
979:
952:
854:
778:
672:
519:
482:
478:
419:
366:
334:
280:
193:
1950:
1914:
1532:
1382:
2409:
2389:
1757:"Mammary Gland Growth Factors: Roles in Normal Development and in Cancer"
1262:
667:
590:
586:
466:
399:
167:
153:
111:
95:
72:
1302:
2653:
2353:
2306:
2086:
2076:
1524:
1060:
546:
498:
457:
443:
375:
325:
308:
215:
which switch development on and off at precise times and places. Here,
99:
1981:
1556:"The organelle of differentiation in embryos: The cell state splitter"
204:
2604:
2572:
2339:
2245:
2013:
Turing, Alan
Mathison (1952). "The chemical basis of morphogenesis".
628:
462:
284:
244:
115:
1478:
2741:
2567:
2536:
2156:
1722:
1422:
1110:
531:
403:
391:
371:
313:
216:
91:
64:
2199:
750:
30:
This article is about the biological process. For other uses, see
2299:
2108:
1614:
486:
395:
149:
251:. An important class of molecules involved in morphogenesis are
67:
to develop its shape. It is one of three fundamental aspects of
2279:
1091:
542:
439:
422:
with the outside. Integrins also serve as receptors to trigger
297:
221:
107:
27:
Biological process that causes an organism to develop its shape
1368:
1150:
890:
Thompson, D'Arcy
Wentworth (2004) , Bonner, John Tyler (ed.),
255:
proteins that determine the fate of cells by interacting with
565:
489:
and the model was later generalized to all of morphogenesis.
349:
events during which epithelial cells become mesenchymal (see
181:
225:, setting up stripes which create the body's segmental form.
2577:
1642:(5th ed.). Oxford University Press. pp. 499–500.
569:
508:
260:
243:
Several types of molecules are important in morphogenesis.
1663:
Miura, T (2008). "Modeling Lung
Branching Morphogenesis".
1287:"Finite element modeling of shape changes in plant cells"
317:
256:
185:
2149:
1333:
1220:(6th ed.). Sunderland, Mass: Sinauer Associates.
2730:
1844:"Cancer morphogenesis: role of mitochondrial failure"
345:. Tissue separation can also occur via more dramatic
51:
creation, literally "the generation of form") is the
1818:. Lippincott Williams & Wilkins. pp. 94–.
1042:
1213:
568:, the morphogenetic proteins encoded by the phage
219:in the fruit fly are switched on by genes such as
2179:. Cambridge, England: Cambridge University Press.
2016:Philosophical Transactions of the Royal Society B
1805:
1285:Bidhendi, Amir J; Geitmann, Anja (January 2018).
1241:Bidhendi, Amir J; Geitmann, Anja (January 2016).
1207:
958:Philosophical Transactions of the Royal Society B
920:
511:a bronchus branches into bronchioles forming the
136:Evolutionary developmental biology § History
2754:
2239:
1284:
1240:
929:On Growth and Form: The Complete Revised Edition
275:, or, more generally, their properties, such as
2183:
1837:
1835:
1748:
1703:
1403:
883:
518:Branching morphogenesis is also evident in the
2174:
1510:
993:
259:. These can be coded for by master regulatory
199:
2225:
1459:
1455:
1453:
1451:
1966:"Structure and function of bacteriophage T4"
1832:
1665:Multiscale Modeling of Developmental Systems
1605:Gordon, Natalie K.; Gordon, Richard (2016).
1604:
1554:Gordon, Natalie K.; Gordon, Richard (2016).
1553:
1278:
1234:
1098:Progress in Biophysics and Molecular Biology
2057:
1964:Yap, Moh Lan; Rossmann, Michael G. (2014).
1963:
1852:Annals of Clinical & Laboratory Science
1754:
785:
279:or cell contractility. For example, during
2232:
2218:
2205:Turing's theory of morphogenesis validated
1761:Cold Spring Harbor Perspectives in Biology
1448:
994:Hiscock, Tom W.; Megason, Sean G. (2015).
806:
492:
2155:
2126:
2085:
2075:
1989:
1788:
1731:
1721:
1581:
1571:
1560:Theoretical Biology and Medical Modelling
1487:
1477:
1431:
1421:
1310:
1261:
1184:
1127:
1109:
1068:
1019:
978:
768:
711:Bioelectricity#Role in pattern regulation
320:(red) or DiO (green). The red cells were
1841:
926:
889:
791:
356:
307:
263:, and either activate or deactivate the
203:
1927:
1637:
1631:
812:
537:
381:
14:
2755:
2722:Index of evolutionary biology articles
2012:
950:
600:Another famous model is the so-called
581:An approach to model morphogenesis in
556:
2213:
2058:Sharpe, James; Green, Jeremy (2015).
1892:
1662:
1656:
953:"The Chemical Basis of Morphogenesis"
944:
526:. Primitive duct formation begins in
433:
1755:Hynes, N. E.; Watson, C. J. (2010).
1704:Fata JE, Werb Z, Bissell MJ (2004).
1404:Fata JE, Werb Z, Bissell MJ (2004).
896:, Cambridge, England; New York, NY:
927:Thompson, D'Arcy Wentworth (1992),
792:Thompson, D'Arcy Wentworth (1917).
173:The Chemical Basis of Morphogenesis
24:
2773:Evolutionary developmental biology
2532:Evolutionary developmental biology
2168:
2111:"Growing Neural Cellular Automata"
1043:Kouros-Mehr, H.; Werb, Z. (2006).
576:
231:evolutionary developmental biology
25:
2784:
2193:
1210:"Morphogenesis and Cell Adhesion"
351:Epithelial–mesenchymal transition
303:
2740:
2489:Evolution of sexual reproduction
1120:10.1016/j.pbiomolbio.2016.08.004
343:differential adhesion hypothesis
188:in 1953, and the development of
2186:Essential Developmental Biology
2143:
2102:
2051:
2006:
1957:
1921:
1886:
1697:
1598:
1547:
1504:
1397:
1362:
1327:
1201:
1144:
1085:
706:Madin-Darby Canine Kidney cells
475:embryonic differentiation waves
316:. Live cells were stained with
312:Cell sorting out with cultured
2260:Genotype–phenotype distinction
1250:Journal of Experimental Botany
1036:
987:
744:
724:
32:Morphogenesis (disambiguation)
13:
1:
2517:Regulation of gene expression
1673:10.1016/S0070-2153(07)81010-6
737:
324:and express higher levels of
314:P19 embryonal carcinoma cells
2687:Endless Forms Most Beautiful
2467:Evolution of genetic systems
2275:Gene–environment correlation
2270:Gene–environment interaction
1943:10.1016/0042-6822(68)90285-7
1907:10.1016/0022-2836(70)90303-7
1895:Journal of Molecular Biology
1348:10.1016/j.biocel.2008.09.027
770:10.1016/j.celrep.2019.07.006
604:, developed in the sixties.
7:
2666:Christiane NĂĽsslein-Volhard
1773:10.1101/cshperspect.a003186
688:Polycystic kidney disease 2
635:
209:Morphogenesis is controlled
200:Genetic and molecular basis
10:
2789:
2542:Hedgehog signaling pathway
2419:Developmental architecture
2188:. Oxford: Wiley-Blackwell.
1336:Int. J. Biochem. Cell Biol
1208:Gilbert, Scott F. (2000).
1012:10.1016/j.cels.2015.12.001
898:Cambridge University Press
800:Cambridge University Press
643:Bone morphogenetic protein
507:In the development of the
496:
228:
133:
129:
71:along with the control of
29:
2719:
2698:
2627:
2555:
2509:
2502:
2466:
2418:
2382:
2369:Transgressive segregation
2315:
2252:
1640:Principles of development
1573:10.1186/s12976-016-0037-2
1169:10.1021/acssynbio.0c00318
663:Reaction–diffusion system
648:Collective cell migration
158:D'Arcy Wentworth Thompson
106:of tissues after damage.
2184:Slack, J. M. W. (2013).
717:
563:bacteriophage (phage) T4
347:cellular differentiation
269:gene regulatory networks
81:cellular differentiation
2547:Notch signaling pathway
2522:Gene regulatory network
2405:Dual inheritance theory
2175:Bard, J. B. L. (1990).
1638:Wolpert, Lewis (2015).
1607:Embryogenesis Explained
931:, New York, NY: Dover,
847:10.1126/science.1164073
701:Cytoplasmic determinant
561:During assembly of the
499:Lung § Development
493:Branching morphogenesis
363:cell adhesion molecules
239:gene regulatory network
2595:cis-regulatory element
2503:Control of development
2383:Non-genetic influences
2349:evolutionary landscape
2128:10.23915/distill.00023
2037:10.1098/rstb.1952.0012
1815:Diseases of the Breast
1460:Sternlicht MD (2006).
980:10.1098/rstb.1952.0012
951:Turing, A. M. (1952).
452:Caenorhabditis elegans
329:
226:
2763:Developmental biology
2706:Nature versus nurture
2610:Cell surface receptor
2527:Evo-devo gene toolkit
2426:Developmental biology
2364:Polygenic inheritance
2290:Quantitative genetics
1383:10.1242/jcs.114.4.629
1216:Developmental biology
1157:ACS Synthetic Biology
653:Embryonic development
497:Further information:
481:morphogenesis during
357:Cell-to-cell adhesion
311:
229:Further information:
207:
134:Further information:
96:maintenance of tissue
88:embryonic development
69:developmental biology
2768:Morphology (biology)
2615:Transcription factor
2330:Genetic assimilation
2317:Genetic architecture
609:computer performance
538:Cancer morphogenesis
388:extracellular matrix
382:Extracellular matrix
253:transcription factor
235:transcription factor
2711:Morphogenetic field
2628:Influential figures
2029:1952RSPTB.237...37T
1970:Future Microbiology
1842:Fosslien E (2008).
1303:10.1104/pp.17.01684
971:1952RSPTB.237...37T
872:on 28 November 2014
839:2008Sci...322.1502M
833:(5907): 1502–1505,
816:(5 December 2008),
557:Virus morphogenesis
424:signal transduction
322:genetically altered
2400:Genomic imprinting
2077:10.1242/dev.114991
1525:10.1007/BF02797122
1263:10.1093/jxb/erv535
1061:10.1002/dvdy.20978
893:On Growth and Form
814:Montell, Denise J.
795:On Growth and Form
613:cellular automaton
503:Breast development
434:Cell contractility
430:ductal branching.
410:-binding proteins
330:
227:
213:"toolkit" of genes
163:On Growth and Form
156:, were written by
53:biological process
2728:
2727:
2661:Eric F. Wieschaus
2623:
2622:
2441:Pattern formation
2345:Fitness landscape
1982:10.2217/fmb.14.91
1976:(12): 1319–1327.
1825:978-1-4511-4870-1
1710:Breast Cancer Res
1649:978-0-19-967814-3
1624:978-981-4350-48-8
1466:Breast Cancer Res
1410:Breast Cancer Res
1227:978-0-87893-243-6
938:978-0-486-67135-2
907:978-0-521-43776-9
658:Pattern formation
602:French flag model
589:can be traced to
339:Malcolm Steinberg
190:molecular biology
160:in his 1917 book
16:(Redirected from
2780:
2745:
2744:
2736:
2671:William McGinnis
2640:Richard Lewontin
2635:C. H. Waddington
2507:
2506:
2484:Neutral networks
2234:
2227:
2220:
2211:
2210:
2189:
2180:
2162:
2161:
2159:
2147:
2141:
2140:
2130:
2106:
2100:
2099:
2089:
2079:
2070:(7): 1203–1211.
2055:
2049:
2048:
2010:
2004:
2003:
1993:
1961:
1955:
1954:
1925:
1919:
1918:
1890:
1884:
1883:
1881:
1875:. Archived from
1848:
1839:
1830:
1829:
1809:
1803:
1802:
1792:
1752:
1746:
1745:
1735:
1725:
1701:
1695:
1694:
1660:
1654:
1653:
1635:
1629:
1628:
1602:
1596:
1595:
1585:
1575:
1551:
1545:
1544:
1508:
1502:
1501:
1491:
1481:
1457:
1446:
1445:
1435:
1425:
1401:
1395:
1394:
1377:(Pt 4): 629–41.
1366:
1360:
1359:
1331:
1325:
1324:
1314:
1291:Plant Physiology
1282:
1276:
1275:
1265:
1247:
1238:
1232:
1231:
1219:
1205:
1199:
1198:
1188:
1148:
1142:
1141:
1131:
1113:
1089:
1083:
1082:
1072:
1040:
1034:
1033:
1023:
991:
985:
984:
982:
948:
942:
941:
924:
918:
917:
916:
914:
887:
881:
880:
879:
877:
871:
865:, archived from
822:
810:
804:
803:
789:
783:
782:
772:
763:(5): 1237–1250.
748:
731:
728:
621:machine learning
617:gradient descent
607:Improvements in
583:computer science
520:ductal formation
513:respiratory tree
146:natural patterns
124:dysmorphogenesis
21:
2788:
2787:
2783:
2782:
2781:
2779:
2778:
2777:
2753:
2752:
2751:
2739:
2731:
2729:
2724:
2715:
2694:
2681:Sean B. Carroll
2619:
2551:
2498:
2462:
2414:
2395:Maternal effect
2378:
2311:
2248:
2238:
2196:
2171:
2169:Further reading
2166:
2165:
2148:
2144:
2107:
2103:
2056:
2052:
2011:
2007:
1962:
1958:
1926:
1922:
1891:
1887:
1879:
1846:
1840:
1833:
1826:
1810:
1806:
1753:
1749:
1702:
1698:
1683:
1661:
1657:
1650:
1636:
1632:
1625:
1603:
1599:
1552:
1548:
1513:Cell Biophysics
1509:
1505:
1479:10.1186/bcr1368
1458:
1449:
1402:
1398:
1367:
1363:
1332:
1328:
1283:
1279:
1245:
1239:
1235:
1228:
1206:
1202:
1149:
1145:
1090:
1086:
1055:(12): 3404–12.
1041:
1037:
992:
988:
949:
945:
939:
925:
921:
912:
910:
908:
888:
884:
875:
873:
869:
820:
811:
807:
790:
786:
749:
745:
740:
735:
734:
729:
725:
720:
715:
683:Eye development
638:
579:
577:Computer models
559:
540:
505:
495:
448:model organisms
436:
384:
359:
306:
241:
202:
142:
132:
35:
28:
23:
22:
15:
12:
11:
5:
2786:
2776:
2775:
2770:
2765:
2750:
2749:
2726:
2725:
2720:
2717:
2716:
2714:
2713:
2708:
2702:
2700:
2696:
2695:
2693:
2692:
2691:
2690:
2678:
2673:
2668:
2663:
2658:
2657:
2656:
2645:François Jacob
2642:
2637:
2631:
2629:
2625:
2624:
2621:
2620:
2618:
2617:
2612:
2607:
2602:
2597:
2592:
2587:
2582:
2581:
2580:
2570:
2565:
2559:
2557:
2553:
2552:
2550:
2549:
2544:
2539:
2534:
2529:
2524:
2519:
2513:
2511:
2504:
2500:
2499:
2497:
2496:
2491:
2486:
2481:
2476:
2470:
2468:
2464:
2463:
2461:
2460:
2455:
2450:
2445:
2444:
2443:
2438:
2428:
2422:
2420:
2416:
2415:
2413:
2412:
2407:
2402:
2397:
2392:
2386:
2384:
2380:
2379:
2377:
2376:
2374:Sequence space
2371:
2366:
2361:
2356:
2351:
2342:
2337:
2332:
2327:
2321:
2319:
2313:
2312:
2310:
2309:
2304:
2303:
2302:
2292:
2287:
2282:
2277:
2272:
2267:
2262:
2256:
2254:
2250:
2249:
2237:
2236:
2229:
2222:
2214:
2208:
2207:
2202:
2195:
2194:External links
2192:
2191:
2190:
2181:
2170:
2167:
2164:
2163:
2142:
2101:
2050:
2023:(641): 37–72.
2005:
1956:
1937:(4): 550–563.
1920:
1901:(3): 293–306.
1885:
1882:on 2017-09-21.
1859:(4): 307–329.
1831:
1824:
1804:
1767:(8): a003186.
1747:
1723:10.1186/bcr634
1696:
1681:
1655:
1648:
1630:
1623:
1597:
1546:
1503:
1447:
1423:10.1186/bcr634
1396:
1361:
1326:
1277:
1256:(2): 449–461.
1233:
1226:
1200:
1163:(2): 277–285.
1143:
1084:
1035:
1006:(6): 408–416.
986:
965:(641): 37–72.
943:
937:
919:
906:
882:
805:
784:
742:
741:
739:
736:
733:
732:
722:
721:
719:
716:
714:
713:
708:
703:
698:
690:
685:
680:
675:
670:
665:
660:
655:
650:
645:
639:
637:
634:
595:Turing pattern
578:
575:
558:
555:
539:
536:
494:
491:
471:microfilaments
435:
432:
383:
380:
358:
355:
305:
304:Cellular basis
302:
273:cell migration
201:
198:
140:Turing pattern
131:
128:
118:of new forms.
55:that causes a
26:
9:
6:
4:
3:
2:
2785:
2774:
2771:
2769:
2766:
2764:
2761:
2760:
2758:
2748:
2743:
2738:
2737:
2734:
2723:
2718:
2712:
2709:
2707:
2704:
2703:
2701:
2697:
2689:
2688:
2684:
2683:
2682:
2679:
2677:
2674:
2672:
2669:
2667:
2664:
2662:
2659:
2655:
2652:
2651:
2650:
2649:Jacques Monod
2646:
2643:
2641:
2638:
2636:
2633:
2632:
2630:
2626:
2616:
2613:
2611:
2608:
2606:
2603:
2601:
2598:
2596:
2593:
2591:
2588:
2586:
2583:
2579:
2576:
2575:
2574:
2571:
2569:
2566:
2564:
2563:Homeotic gene
2561:
2560:
2558:
2554:
2548:
2545:
2543:
2540:
2538:
2535:
2533:
2530:
2528:
2525:
2523:
2520:
2518:
2515:
2514:
2512:
2508:
2505:
2501:
2495:
2492:
2490:
2487:
2485:
2482:
2480:
2477:
2475:
2472:
2471:
2469:
2465:
2459:
2456:
2454:
2451:
2449:
2446:
2442:
2439:
2437:
2434:
2433:
2432:
2431:Morphogenesis
2429:
2427:
2424:
2423:
2421:
2417:
2411:
2408:
2406:
2403:
2401:
2398:
2396:
2393:
2391:
2388:
2387:
2385:
2381:
2375:
2372:
2370:
2367:
2365:
2362:
2360:
2357:
2355:
2352:
2350:
2346:
2343:
2341:
2338:
2336:
2333:
2331:
2328:
2326:
2323:
2322:
2320:
2318:
2314:
2308:
2305:
2301:
2298:
2297:
2296:
2293:
2291:
2288:
2286:
2283:
2281:
2278:
2276:
2273:
2271:
2268:
2266:
2265:Reaction norm
2263:
2261:
2258:
2257:
2255:
2251:
2247:
2243:
2235:
2230:
2228:
2223:
2221:
2216:
2215:
2212:
2206:
2203:
2201:
2198:
2197:
2187:
2182:
2178:
2173:
2172:
2158:
2153:
2146:
2138:
2134:
2129:
2124:
2120:
2116:
2112:
2105:
2097:
2093:
2088:
2083:
2078:
2073:
2069:
2065:
2061:
2054:
2046:
2042:
2038:
2034:
2030:
2026:
2022:
2018:
2017:
2009:
2001:
1997:
1992:
1987:
1983:
1979:
1975:
1971:
1967:
1960:
1952:
1948:
1944:
1940:
1936:
1932:
1924:
1916:
1912:
1908:
1904:
1900:
1896:
1889:
1878:
1874:
1870:
1866:
1862:
1858:
1854:
1853:
1845:
1838:
1836:
1827:
1821:
1817:
1816:
1808:
1800:
1796:
1791:
1786:
1782:
1778:
1774:
1770:
1766:
1762:
1758:
1751:
1743:
1739:
1734:
1729:
1724:
1719:
1715:
1711:
1707:
1700:
1692:
1688:
1684:
1682:9780123742537
1678:
1674:
1670:
1666:
1659:
1651:
1645:
1641:
1634:
1626:
1620:
1616:
1612:
1608:
1601:
1593:
1589:
1584:
1579:
1574:
1569:
1565:
1561:
1557:
1550:
1542:
1538:
1534:
1530:
1526:
1522:
1518:
1514:
1507:
1499:
1495:
1490:
1485:
1480:
1475:
1471:
1467:
1463:
1456:
1454:
1452:
1443:
1439:
1434:
1429:
1424:
1419:
1415:
1411:
1407:
1400:
1392:
1388:
1384:
1380:
1376:
1372:
1365:
1357:
1353:
1349:
1345:
1342:(2): 349–69.
1341:
1337:
1330:
1322:
1318:
1313:
1308:
1304:
1300:
1296:
1292:
1288:
1281:
1273:
1269:
1264:
1259:
1255:
1251:
1244:
1237:
1229:
1223:
1218:
1217:
1211:
1204:
1196:
1192:
1187:
1182:
1178:
1174:
1170:
1166:
1162:
1158:
1154:
1147:
1139:
1135:
1130:
1125:
1121:
1117:
1112:
1107:
1103:
1099:
1095:
1088:
1080:
1076:
1071:
1066:
1062:
1058:
1054:
1050:
1046:
1039:
1031:
1027:
1022:
1017:
1013:
1009:
1005:
1001:
997:
990:
981:
976:
972:
968:
964:
960:
959:
954:
947:
940:
934:
930:
923:
909:
903:
899:
895:
894:
886:
868:
864:
860:
856:
852:
848:
844:
840:
836:
832:
828:
827:
819:
815:
809:
801:
797:
796:
788:
780:
776:
771:
766:
762:
758:
754:
747:
743:
727:
723:
712:
709:
707:
704:
702:
699:
697:
696:embryogenesis
695:
691:
689:
686:
684:
681:
679:
678:Axon guidance
676:
674:
671:
669:
666:
664:
661:
659:
656:
654:
651:
649:
646:
644:
641:
640:
633:
631:
630:
624:
622:
618:
614:
610:
605:
603:
598:
596:
592:
588:
584:
574:
571:
567:
564:
554:
552:
551:Mitochondrial
548:
544:
535:
533:
529:
525:
524:mammary gland
521:
516:
514:
510:
504:
500:
490:
488:
484:
480:
476:
472:
468:
464:
460:
459:
454:
453:
449:
445:
441:
431:
429:
428:mammary gland
425:
421:
417:
413:
409:
408:microfilament
405:
401:
397:
393:
389:
379:
377:
373:
368:
364:
354:
352:
348:
344:
340:
336:
327:
323:
319:
315:
310:
301:
299:
295:
291:
286:
282:
278:
277:cell adhesion
274:
270:
266:
265:transcription
262:
258:
254:
250:
246:
240:
236:
232:
224:
223:
218:
214:
210:
206:
197:
195:
191:
187:
183:
179:
175:
174:
169:
165:
164:
159:
155:
151:
147:
141:
137:
127:
125:
119:
117:
113:
109:
105:
101:
97:
93:
89:
84:
82:
78:
74:
73:tissue growth
70:
66:
62:
58:
54:
50:
46:
43:
39:
38:Morphogenesis
33:
19:
18:Morphogenetic
2685:
2578:eyeless gene
2474:Evolvability
2448:Segmentation
2430:
2325:Canalisation
2295:Heterochrony
2285:Heritability
2253:Key concepts
2241:
2185:
2176:
2145:
2118:
2114:
2104:
2067:
2063:
2053:
2020:
2014:
2008:
1973:
1969:
1959:
1934:
1930:
1923:
1898:
1894:
1888:
1877:the original
1856:
1850:
1814:
1807:
1764:
1760:
1750:
1713:
1709:
1699:
1664:
1658:
1639:
1633:
1615:10.1142/8152
1606:
1600:
1563:
1559:
1549:
1516:
1512:
1506:
1469:
1465:
1413:
1409:
1399:
1374:
1370:
1364:
1339:
1335:
1329:
1297:(1): 41–56.
1294:
1290:
1280:
1253:
1249:
1236:
1215:
1203:
1160:
1156:
1146:
1104:(1): 58–69.
1101:
1097:
1087:
1052:
1048:
1038:
1003:
1000:Cell Systems
999:
989:
962:
956:
946:
928:
922:
911:, retrieved
892:
885:
874:, retrieved
867:the original
830:
824:
808:
794:
787:
760:
757:Cell Reports
756:
746:
726:
693:
673:Gastrulation
627:
625:
606:
599:
580:
560:
541:
517:
506:
483:gastrulation
479:neural plate
467:microtubules
456:
450:
437:
420:cytoskeleton
418:to link the
385:
367:cell culture
360:
341:through his
335:cell sorting
331:
283:, clumps of
281:gastrulation
242:
220:
194:biochemistry
178:spiral shell
171:
161:
148:such as the
143:
120:
104:regeneration
85:
48:
44:
37:
36:
2676:Mike Levine
2585:Distal-less
2410:Polyphenism
2390:Epigenetics
2242:development
2087:10230/25028
2064:Development
1716:(1): 1–11.
1519:: 177–238.
1416:(1): 1–11.
913:11 December
876:11 December
668:Neurulation
591:Alan Turing
587:mathematics
528:development
444:germ layers
400:fibronectin
168:Alan Turing
154:phyllotaxis
112:unicellular
2757:Categories
2654:Lac operon
2479:Robustness
2458:Modularity
2453:Metamerism
2359:Plasticity
2354:Pleiotropy
2307:Heterotopy
2157:2103.08737
1472:(1): 201.
1371:J Cell Sci
1111:1702.03337
738:References
694:Drosophila
547:metastasis
458:Drosophila
376:E-cadherin
326:E-cadherin
285:stem cells
245:Morphogens
100:stem cells
77:patterning
47:shape and
40:(from the
2605:Morphogen
2590:Engrailed
2573:Pax genes
2494:Tinkering
2340:Epistasis
2335:Dominance
2246:phenotype
2137:213719058
1781:1943-0264
1177:2161-5063
629:Minecraft
463:zebrafish
412:α-actinin
404:integrins
372:cadherins
249:receptors
217:gap genes
116:evolution
2568:Hox gene
2556:Elements
2537:Homeobox
2096:25804733
2000:25517898
1931:Virology
1865:18988924
1799:20554705
1742:14680479
1691:18023732
1592:26965444
1498:16524451
1442:14680479
1391:11171368
1356:18848899
1321:29229695
1272:26689854
1195:33449631
1138:27544910
1079:17039550
1049:Dev. Dyn
1030:26771020
863:27982230
855:19056976
779:31365867
636:See also
532:estrogen
392:Collagen
294:Hedgehog
92:organism
65:organism
2747:Biology
2699:Debates
2510:Systems
2436:Eyespot
2300:Neoteny
2115:Distill
2025:Bibcode
1991:4275845
1951:4878023
1915:4907266
1873:4538888
1790:2908768
1583:4785624
1541:4349055
1533:2450659
1489:1413974
1312:5761827
1186:8486170
1129:5563449
1070:2730892
1021:4707970
967:Bibcode
835:Bibcode
826:Science
522:of the
487:axolotl
485:of the
446:in the
396:laminin
298:ephrins
170:in his
150:spirals
130:History
49:genesis
2733:Portal
2600:Ligand
2280:Operon
2135:
2094:
2045:937133
2043:
1998:
1988:
1949:
1913:
1871:
1863:
1822:
1797:
1787:
1779:
1740:
1733:314442
1730:
1689:
1679:
1646:
1621:
1590:
1580:
1566:: 11.
1539:
1531:
1496:
1486:
1440:
1433:314442
1430:
1389:
1354:
1319:
1309:
1270:
1224:
1193:
1183:
1175:
1136:
1126:
1077:
1067:
1028:
1018:
935:
904:
861:
853:
777:
566:virion
543:Cancer
501:, and
440:myosin
398:, and
296:, and
237:, and
222:bicoid
138:, and
108:Cancer
102:or in
90:of an
61:tissue
45:morphĂŞ
2152:arXiv
2133:S2CID
2121:(2).
2041:S2CID
1880:(PDF)
1869:S2CID
1847:(PDF)
1537:S2CID
1246:(PDF)
1106:arXiv
870:(PDF)
859:S2CID
821:(PDF)
718:Notes
570:genes
416:talin
261:genes
211:by a
182:snail
180:of a
42:Greek
2240:The
2092:PMID
1996:PMID
1947:PMID
1911:PMID
1861:PMID
1820:ISBN
1795:PMID
1777:ISSN
1738:PMID
1687:PMID
1677:ISBN
1644:ISBN
1619:ISBN
1588:PMID
1529:PMID
1494:PMID
1438:PMID
1387:PMID
1352:PMID
1317:PMID
1268:PMID
1222:ISBN
1191:PMID
1173:ISSN
1134:PMID
1075:PMID
1026:PMID
933:ISBN
915:2012
902:ISBN
878:2012
851:PMID
775:PMID
509:lung
469:and
461:and
414:and
386:The
192:and
166:and
75:and
57:cell
2244:of
2123:doi
2082:hdl
2072:doi
2068:142
2033:doi
2021:237
1986:PMC
1978:doi
1939:doi
1903:doi
1785:PMC
1769:doi
1728:PMC
1718:doi
1669:doi
1611:doi
1578:PMC
1568:doi
1521:doi
1484:PMC
1474:doi
1428:PMC
1418:doi
1379:doi
1375:114
1344:doi
1307:PMC
1299:doi
1295:176
1258:doi
1181:PMC
1165:doi
1124:PMC
1116:doi
1102:122
1065:PMC
1057:doi
1053:235
1016:PMC
1008:doi
975:doi
963:237
843:doi
831:322
765:doi
585:or
318:DiI
290:Wnt
257:DNA
186:DNA
152:of
98:by
79:of
63:or
2759::
2647:+
2131:.
2117:.
2113:.
2090:.
2080:.
2066:.
2062:.
2039:.
2031:.
2019:.
1994:.
1984:.
1972:.
1968:.
1945:.
1935:35
1933:.
1909:.
1899:47
1897:.
1867:.
1857:38
1855:.
1849:.
1834:^
1793:.
1783:.
1775:.
1763:.
1759:.
1736:.
1726:.
1712:.
1708:.
1685:.
1675:.
1617:.
1609:.
1586:.
1576:.
1564:13
1562:.
1558:.
1535:.
1527:.
1517:11
1515:.
1492:.
1482:.
1468:.
1464:.
1450:^
1436:.
1426:.
1412:.
1408:.
1385:.
1373:.
1350:.
1340:41
1338:.
1315:.
1305:.
1293:.
1289:.
1266:.
1254:67
1252:.
1248:.
1212:.
1189:.
1179:.
1171:.
1161:10
1159:.
1155:.
1132:.
1122:.
1114:.
1096:.
1073:.
1063:.
1051:.
1047:.
1024:.
1014:.
1002:.
998:.
973:.
961:.
955:.
900:,
857:,
849:,
841:,
829:,
823:,
798:.
773:.
761:28
759:.
755:.
597:.
549:.
455:,
394:,
300:.
292:,
233:,
196:.
126:.
83:.
59:,
2735::
2347:/
2233:e
2226:t
2219:v
2160:.
2154::
2139:.
2125::
2119:5
2098:.
2084::
2074::
2047:.
2035::
2027::
2002:.
1980::
1974:9
1953:.
1941::
1917:.
1905::
1828:.
1801:.
1771::
1765:2
1744:.
1720::
1714:6
1693:.
1671::
1652:.
1627:.
1613::
1594:.
1570::
1543:.
1523::
1500:.
1476::
1470:8
1444:.
1420::
1414:6
1393:.
1381::
1358:.
1346::
1323:.
1301::
1274:.
1260::
1230:.
1197:.
1167::
1140:.
1118::
1108::
1081:.
1059::
1032:.
1010::
1004:1
983:.
977::
969::
845::
837::
802:.
781:.
767::
34:.
20:)
Text is available under the Creative Commons Attribution-ShareAlike License. Additional terms may apply.