Branching morphogenesis – Dr Marcus Fruttiger’s lecture
Branching morphogenesis – (bottom) Jenny E. Sabin
General points
Even though all organisms are 3 dimensional “objects” metabolic exchange processes occur over 2 dimensional surfaces. In single cell organisms the surface of the organism is sufficient for metabolic exchange with the environment. However, in bigger organisms metabolic exchange surfaces have to be much larger. These exchange surfaces are folded and fragmented so they can be packed inside the body of animals (unlike plants). For example, gas exchange in the human lung occurs across the surface of about 300 million alveoli with a total surface of about 100 m2; nutrient and gas exchange in the human vasculature takes place across the total surface of about 600 m2. Evolution has favored a branched morphology of such metabolically active surfaces, maximizing the active surface and minimizing transport distances. Lungs, kidneys, mammary glands, salivary glands or the vasculature all display a branched morphology.
How to build branched structures
In principle, a branched structure can be built by the iterative use of a few simple subroutines. These are the initiation of a bud, the extension of a bud and the splitting at the end of a bud. With this simple set of morphogenetic units complex structures can be built. The formation of new buds, creating new centers of growth, is important for the overall morphology of the branched structure. For example in the human lung different modes of bud formation occur during different stages of development. In the early phases of bronchi formation new buds arise at the side of an extended bud, a process termed ‘secondary budding’. At later stages new buds form at the end of an extended bud (‘branching’) and in the final stages during alveoli formation terminal sacs are split in several compartments by invagination (‘splitting’).
Drosophila tracheal system
The tracheal system in drosophila larvae is a relatively simple model system for the study of branched structures and has provided some amazing insights into the biology of branching morphology. The system is a tubular network formed by a monolayer epithelium (Fig.1). Oxygen enters through the spiracular openings and diffuses freely to the target tissues. The network has three branching levels: primary branches, secondary branches and blind ending terminal branches. The first two branching levels display a stereotypical morphology whereas the terminal branches are not stereotypical. Terminal branches are very thin cytoplasmic extensions contacting in many tissues almost every single cell. The reproducible pattern of the primary and secondary branches implies a tight morphogenetic program responsible for the development of the network.
Morphogenetic control of the drosophila tracheal system
Genetic screens of drosophila larvae have revealed more than 50 genes involved in the formation of the tracheal system. Some of the genes are used during each generation of branch formation whereas others are specifically involved in early or late stages of the developing network. A gene expressed very early in development is trachealess, a transcription factor, which appears in 10 patches bilaterally along the longitudinal axes of the larvae. This initiates tissue locally to form slight invaginations (sacs) and defines cells as future components of the tracheal system. Without it no trachea will form (hence the name). It also induces the expression of breathless in these sacs, a drosophila ortholog of the mammalian fibroblast growth factor receptor (FGFR). The ligand for this receptor is branchless (drosophila ortholog of FGF) and is dynamically expressed in 6 patches of mesenchymal tissue around each sac. This diffusible factor induces bud formation and bud extension from the sacs resulting in 6 primary branches. Some of these branches fuse forming a longitudinal continues tube (involving a gene called escargot). The other branches grow closer to the branchless expressing centers exposing the tip of the growing branch to high concentrations of Branchless. This high concentration of Branchless induces the expression of pointed (transcription factor) and sprouty (antagonist of branchless) at the tip of the branch. Pointed causes the tip of the branch split (secondary branches) and sprouty restricts branching to the tip by inhibiting branching further back along the extended branch. Secondary branches also express pruned (transcription factor) which is a prerequisite for the formation of terminal branches. These terminal branches are very thin cellular processes growing towards cells secreting Branchless. However, this time branchless is not expressed stereotypical but is induced in any cell experiencing hypoxia. In summary, the diffusible factor Branchless is used three times in the formation of the drosophila tracheal system but at each level in a different context. At the level of primary branches branchless induces bud formation and extension, at the secondary level high concentrations of Branchless induces the expression of genes involved in secondary branching and at the level of terminal branches branchless mediates the need of oxygen-starved cells for oxygen by promoting and directing the growth of terminal branches.
Mammalian lung
Like most internal organs the lungs develop from the foregut (in the early embryo a ventral longitudinal tube). Initially, two buds extend from the foregut resulting in the left and right bronchus. In the mouse four secondary buds extend from the two initial branches (three on the right-hand side and one on the left) giving rise to four lung lobes. The genetic control of these processes is not completely understood but it has been shown that Gli genes are involved in the induction of the secondary buds. It is surprising that also FGF (human ortholog to drosophila branchless) is crucially involved in mammalian lung formation. Mammals possess more than one FGF gene. Currently there are more than 20 different FGFs known and many of them have overlapping functions. However, “knock out mice” lacking FGF10 are born without lungs and limbs (limbs are also formed from buds!). The function of FGF in lung development is similar to branchless in drosophila trachea formation. The underlying principle is again a mesenchymal-epithelial cell-cell interaction mediated by FGF. Epithelial cells, expressing FGF receptor, respond to the secretion of FGF from nearby mesenchyme by bud formation and bud extension towards the FGF source. Exposure of the branch tip to high concentrations of FGF induces the expression of secondary genes in the tip such as bone morphogenetic protein 4 (BMP4), sonic hedgehog (Shh) and a mammalian sprouty ortholog (Sprouty 2), thus, turning the tips of the bronchial branches into signaling centers. BMP4 inhibits epithelial cell proliferation limiting branch extension. Shh is proposed to inhibit FGF10 expression in the mesenchyme near the tip, which splits FGF10 expression promoting the next round of branching and Sprouty2 (like drosophila sprouty) restricts branching to the tip of the branch.
Summary
In order to form a branching structure an iterative process of bud formation, bud extension and branching is required. It is remarkable that FGF-FGFR interactions appear to be a central unit repetitively used to generate the branching morphology in both the drosophila tracheal system and the mammalian lung.
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Plus t? “Donc, le ministre du redressement productif lance aux jeunes ingénieurs “Nous voulons les en allant à leur rencontre explique son cabinet Ils ont un rle essentiel à dans la réindustrialisation du pays car il n’y a pas d’ sans ingénieurs” Or ces forces vives ont souvent une mauvaise image du secteur et s’en détournent préférant le col blanc au col bleu Le ministre saura-t-il les Le Mondefr a le plaisir de vous la lecture de cet article habituellement réservé aux abonnés du MondefrProfitez de tous les articles réservés du Mondefr en Voir l’infographie : M Montebourg intervient dans un contexte favorable La dernière de l’une des plus prestigieuses écoles d’ingénieurs du pays Polytechnique montre un regain d’intérêt pour l’industrie Parmi les diplmés 2011 qui débutent leur carrière en entreprise (soit la moitié de la promotion) 465 % ont choisi l’industrie Ils n’étaient que 37 % l’année précédente Ceux qui optent pour la banque-assurance n’étaient plus que 13 % en 2011 contre 28 % en 2010 Il faut cependant le nombre de diplmés qui vont dans le conseil : 22 % en 2011 contre 17 % en 2010″ILS SONT TENTS D’ VERS CE QUI BRILLE”Ces chiffres rejoignent ceux du Conseil national des ingénieurs et scientifiques de France (CNISF) Dans sa dernière enquête annuelle (qui donne une photographie des 750 000 ingénieurs franais) le CNISF indique qu’entre fin 2010 et fin 2011 le pourcentage de débutants rejoignant l’industrie est passé de 42 % à 46 % ce qui montre bien que s’il n’y a pas d’industrie sans ingénieurs il y a beaucoup d’ingénieurs hors industrie Et certains d’entre eux se dirigent notamment vers un secteur très symbolique : “Les nouveaux diplmés note le CNISF ont été de plus en plus nombreux à leur carrière dans les secteurs de la finance et du conseil durant les quinze dernières années et ils y sont restés”Lire aussi : Julien Roitman président du CNISF se souvient de sa surprise lorsqu’il a assisté il y a quelques années à Londres à la création de l’association locale des élec : “Sur les 60 ingénieurs présents 50 travaillaient dans la finance Et ce n’est pas un cas isolé” “C’est sr qu’ils sont tentés d’ vers ce qui brille” reconnat Laurent Billès-Garabédian président de l’Association des anciens de polytechnique l’AX Et pour des jeunes aussi prometteurs rien n’est trop beau “Il y a quelques années raconte le président de l’AX des banques anglo-saxonnes n’hésitaient pas pour de jeunes polytechniciens à les à dner au Crillon Le recruteur leur tenait un discours du genre je suis venu de Londres en jet privé uniquement pour vous. C’est juste nul Mais a impressionne Bon la crise est passée là-dessus”ARGUMENT MORALEt ce sont dans les meilleures écoles que la tentation est la plus forte Toujours selon le CNISF 19 % des diplmés de moins de 30 ans des sept meilleures écoles d’ingénieurs ont fait le choix de la finance et du conseil (103 % pour la finance uniquement) Chez les “autres” la proportion n’est que de 6 % (17 % pour la seule finance) Les données de Polytechnique sont d’autant plus intéressantes Elles montrent peut-être que la déroute industrielle du pays devenue patente un déclic s’est produit chez les ingénieurs les plus réputésMais l’appel de M Montebourg s’adresse à tous les ingénieurs Comme l’argument moral “Nos étudiants payent 800 euros une année d’études qui revient à 15 000 euros à la nation rappelle tout nouveau directeur de l’Ecole des mines d’Alès Ils ont donc un moral vis-à-vis de leur pays celui de créer de la valeur Et cela passe par l’industrie Nous avons pour mission de la développer Et il faut en être “Les mentalités devront évoluer en profondeur reconnat M Goubert : “Les écoles ont contribué en ne le combattant pas assez à l’installation du discours qui a dominé les vingt dernières années du XXe siècle Qu’entendait-on alors L’industrie c’est sale L’industrie ce sont les grands sinistres sidérurgiques et charbonniers mieux vaut donc se vers le service Mais aussi : faisons de l’industrie sans usine Voilà comment les écoles d’ingénieurs ont formé une génération de banquiers. Comme leurs élèves avaient un bon niveau scientifique ils sont partis des statistiques dans les banques””DES FRUITS SUR PLUSIEURS ANNES”L’enseignement aussi doit évoluer Pendant longtemps les écoles d’ingénieurs ont formé de futurs salariés notamment de grands groupes “Un phénomène nouveau est en train d’émerger assure La entrepreneuriale est de plus en plus présente dans les enseignements” De fait le redémarrage de l’industrie passe par le développement de l’innovation “Aujourd’hui poursuit M Roitman l’écrasante majorité des start-up sont créées par des diplmés d’écoles de commerce Il n’est donc pas étonnant qu’il s’agisse surtout de Les jeunes ingénieurs s’ils étaient encouragés à créer des PME le feraient plus facilement dans l’industrie Bref ce mouvement pourrait des fruits sur plusieurs années”Surtout note M Goubet le directeur de l’Ecole des mines d’Alès les liens avec l’industrie relèverait du gagnant-gagnant “Je mets mes ingénieurs à la disposition des de la région précise-t-il Ils aident à développer des produits ou à améliorer les processus et ils se forment” M Billès-Garabédian abonde dans son sens : “Il faut davantage d’industriel dans le cursus notamment en faisant des patrons d’industrie” La ministre de l’ Geneviève Fioraso souligne que “l’alternance permet aussi d’ des contacts avec les entreprises Cela pourrait que les étudiants se dirigent plus facilement vers les entreprises industrielles qui innovent Comme ils ne les connaissent pas ils vont vers ce qu’on leur vend : le et la finance”A défaut d’ créé le mouvement (les chiffres de Polytechnique ou du CNISF datent de 2011) M Montebourg peut en accélérer le rythme Mais il n’en recueillera peut-être pas les fruits “Il se passera quatre ou cinq ans avant que des effets importants ou visibles se produisent estime M Roitman Comme dit un proverbe chinois “on entend l’arbre qui tombe pas la forêt qui pousse””ner des poursuites disciplinaires contre un magistrat du parquet”, ne l’a pas fait. 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