Reconstruction d’états ancestraux

Introduction

Si certaines combinaisons de caractères sont systématiquement associées à plusieurs espèces, cela pourrait suggérer que des forces évolutives, comme la sélection, ont façonné ces associations.

Toutefois, les associations non aléatoires de certains traits chez certaines espèces peuvent être dues à l’**héritage commun* de leurs ancêtres et, par conséquent, les changements concomitants dans le temps ne peuvent être déduits.

Si les caractères ont évolué de façon aléatoire sans association, les espèces plus étroitement apparentées sont plus susceptibles d’être semblables que les autres, créant ainsi des relations apparentes entre les caractères.

Il est donc nécessaire de considérer les relations phylogénétiques entre les espèces lors de l’analyse de leurs caractères.

Deux questions peuvent être abordées lors de l’intégration de la phylogénie dans les données comparatives :

prise en compte de la non-indépendance inter-espèces dans l’étude des caractères et de leurs relations,
estimation des paramètres d’évolution des caractères.

Ces deux questions sont étroitement liées. En effet l’impact de la phylogénie sur la distribution des caractères dépend non seulement de la phylogénie mais aussi de la façon dont ces caractères évoluent.

Mise en oeuvre

Vérifiez que l’arbre tr est enraciné

Tracé de l’arbre

Lecture des informations sur les souches

Changement des labels et annotation de l’arbre

Arbre espèce annoté.

GC et taille des génomes

Taux de GC en fonction de la taille des génomes.

Cartographie de l’évolution d’un caractère continue sur l’arbre

La fonction contMap (Map continuous trait evolution on the tree) trace l’arbre sur lequel est reporté l’évolution d’un caractère continu. La cartographie est réalisée en estimant les états aux nœuds internes à l’aide du ML avec la fonction fastAnc, et l’interpolation des états le long de chaque branche (Felsenstein, 1985).

Evolution du taux de GC sur les branches de l’arbre.

Evolution de la taille des génomes sur les branches de l’arbre.

Reconstruction par Maximum de vraisemblance

Il existe plusieurs fonctions R utilisant le ML pour estimer les états ancestraux aux nœuds internes pour les caractères continus:

ace(method=“ML”),
fastAnc,
anc.ML.

Les trois méthodes effectuent l’estimation du ML en supposant un modèle brownien pour modéliser l’évolution du caractère. La seule différence est la méthode d’optimisation.

fastAnc utilise une méthode de ré-enracinement dans laquelle l’arbre est ré-enraciné à chaque nœud interne et l’algorithme de contraste est utilisé pour obtenir l’état ML pour ce noeud.
anc.ML utilise une optimisation numérique, mais initie la recherche avec les valeurs obtenues avec fastAnc.

Comparaison des valeurs estimées aux noeuds par le trois méthodes qui effectuent l’estimation du ML.

Tracé de l’arbre

Etats ancestraux de la fréquence des bases aux noeuds de l’arbre obtenus avec aceML (rouge G+C et bleu A+T).

Lien entre caractères

Felsenstein a proposé une méthode qui prend en compte la phylogénie dans l’analyse des données comparatives (GC contre taille du génome). L’idée qui sous-tend sa méthode des contrastes est que, si nous supposons qu’un trait continu évolue de façon aléatoire dans n’importe quelle direction (modèle de mouvement brownien), le contraste entre deux espèces devrait avoir une distribution normale avec une moyenne nulle et une variance proportionnelle au temps écoulé depuis la divergence. Si les contrastes sont mis à l’échelle avec le temps écoulé depuis la divergence, alors ils ont une variance égale à un.

La fonction pic (phylogenetically independent contrasts)

Méthode des contrastes avec le taux de GC.

Méthode des contrastes avec les tailles de génomes.

Lien entre les pairs de contrastes

Lien entre les pairs de contrastes.

## [1] 0.276208

## 
## Call:
## lm(formula = pic.lns ~ pic.gc)
## 
## Coefficients:
## (Intercept)       pic.gc  
##    -0.01031      0.29176

Remarque, faire la régression entre les PICs en passant par l’origine est justifiée seulement si les caractères évoluent sous un modèle de mouvement brownien et s’il existe une relation linéaire entre eux.

Autocorrélation phylogénétique

Comme les espèces ne sont pas indépendantes par leurs relations phylogénétiques, nous pouvons utiliser cette dépendance pour quantifier l’association entre les variables observées au niveau des espèce. Gittleman et Kot[109] propose une méthode basée sur une approche d’autocorrélation.

Elle utilise l’indice d’autocorrélation I de Moran qui fait intervenir ‘’wij’’ un poids qui quantifie la proximité phylogénétique entre espèces ‘’i’’ et ‘’j’’.

Avec un arbre phylogénétique, nous avons

‘’wij’’ = 1/‘’dij’’,

où ‘’dij’’ sont les distances mesurées sur un arbre, et ‘’wii’’ = 0.

Table: indice d’autocorrélation de Moran pour le taux de GC.
observed	expected	sd	p.value
0.5035547	-0.0666667	0.1456557	9.05e-05

Table: indice d’autocorrélation de Moran pour la taille des génomes.
observed	expected	sd	p.value
0.5037201	-0.0666667	0.1456317	8.98e-05

Le résultat est une liste avec quatre éléments : * la valeur observée de I (observée), * sa valeur attendue sous l’hypothèse nulle d’aucune corrélation, * l’écart-type de l’I (sd) observé, * la P-valeur de l’hypothèse nulle.

La fonction gearymoran d’ade4 calcule le coefficient de Moran et test sa signification à l’aide d’une procédure de randomisation. L’option nrepet spécifie le nombre de réplications de la randomisation (999 par défaut).

## class: krandtest lightkrandtest 
## Monte-Carlo tests
## Call: as.krandtest(sim = matrix(res$result, ncol = nvar, byrow = TRUE), 
##     obs = res$obs, alter = alter, names = test.names)
## 
## Number of tests:   2 
## 
## Adjustment method for multiple comparisons:   none 
## Permutation number:   99999 
##   Test       Obs  Std.Obs     Alter  Pvalue
## 1   gc 0.5930783 3.205909 two-sided 0.00016
## 2   ln 0.5949111 3.211170 two-sided 0.00017

Habitats

Nous allons maintenant nous intéresser à la profondeur à laquelle les organismes vivent. Cette information est absente pour la souche Aaap. Nous allons donc supprimer cette feuille à l’arbre.

Table: Propriétés des souches. Triées selon l’odre des feuilles dans l’arbre espèce.
	Strain	Light	Depth
Aaan	S. CC9605	Syn	51
Aaap	S. WH8102	Syn	NA
Aaam	S. CC9902	Syn	5
Aaao	S. CC9311	Syn	95
Aaac	P. MIT9313	LLIV	135
Aaal	P. MIT9303	LLIV	100
Aaah	P. MIT9211	LLIII	83
Aaaa	P. SS120	LLII	120
Aaad	P. NATL2A	LLI	10
Aaai	P. NATL1A	LLI	30
Aaae	P. MIT9312	HLII	135
Aaak	P. MIT9215	HLII	5
Aaaf	P. AS9601	HLII	50
Aaaj	P. MIT9301	HLII	90
Aaag	P. MIT9515	HLI	15
Aaab	P. MED4	HLI	5

Profondeur à laquelle les organismes vivent.

Ecotypes

Table: Ecotype des souches.
	x
S. CC9605	Syn
S. WH8102	Syn
S. CC9902	Syn
S. CC9311	Syn
P. MIT9313	LL
P. MIT9303	LL
P. MIT9211	LL
P. SS120	LL
P. NATL2A	LL
P. NATL1A	LL
P. MIT9312	HL
P. MIT9215	HL
P. AS9601	HL
P. MIT9301	HL
P. MIT9515	HL
P. MED4	HL

Reconstruction des états du caractère aux nœuds de l’arbre avec la fonction ace d’ape (type=“discrete”, model=“ER” for equal-rates model).

Les vitesses de transition entre les trois états du carctère sont identiques.

Origine des ecotypes.

Origine de tous les ecotypes.

Profils des gènes

Transposer la matrice et ordonner les lignes comme les feuilles de l’arbre

Compiler les différents motifs de présence absence des gènes observés dans chaque groupe de gènes orthologues dans une liste

Définir un ensemble de motifs

Pour la mise en main des scripts, il peut être utile de travailler avec un sous ensemble de motifs!

Taille des patterns trouvés

Occurence des motifs trouvés.

Inférence des états ancestraux

Les différentes fonctions utilisent des paramètres similaires.

Le nombre d’états est déduit des données.

Le modèle est spécifié avec une matrice d’entiers représentant les indices des paramètres : 1 représente le premier paramètre, 2 le second, et ainsi de suite.

Vous devrez choisir la matrice qui spécifie les probabilités de transition entre les états de votre caractère discret. Ace propose des raccourcis pour

un modèle à un paramètre (ER) où tous les taux de transitions sont égaux,
un modèle symétrique (SYM) dans lequel les taux de transitions dans les deux sens sont égaux,
un modèle où tous les taux sont différents (ARD), toutes les transitions possibles entre les états reçoivent des paramètres distincts.

Vous pouvez également spécifier votre propre matrice. Pour indiquer qu’une transition est impossible, un zéro doit être donné dans la cellule appropriée de la matrice.

La reconstruction la plus parcimonieuse

Cette fonction MPR (Most Parsimonious Reconstruction) réalise la reconstruction des caractères ancestraux par parcimonie, comme décrit dans Hanazawa et al. (1995) et modifié par Narushima et Hanazawa (1997). Ils ont utilisé l’approche proposée par Farris (1970) et Swofford and Maddison’s (1987) pour reconstruire les états ancestraux. Le caractère est supposé prendre des valeurs entières.

L’algorithme recherche les ensembles de valeurs pour chaque nœud sous forme d’intervalles avec des valeurs inférieures et supérieures (ex. [1,1]).

L’arbre doit être non enraciné et entièrement dichotomique.

Exemple

Reconstruction des caractères ancestraux par parcimonie.

Tracé MPR pour chaque motif

Reconstruction des caractères ancestraux par parcimonie.

Marquage stochastique des caractères

La fonction make.simmap de phytools permet de réaliser un marquage stochastique des caractères aux nœuds de l’arbre.

nsim: number of simulations.
pi: gives the prior distribution on the root node of the tree, options are equal, estimated, or a vector with the frequencies.

Exemple

## Using pi estimated from the stationary distribution of Q assuming a flat prior.
## pi =
##   0   1 
## 0.5 0.5 
## 
## make.simmap is sampling character histories conditioned on
## the transition matrix
## 
## Q =
##          0        1
## 0 -1.13569  1.13569
## 1  1.13569 -1.13569
## (estimated using likelihood);
## and (mean) root node prior probabilities
## pi =
##   0   1 
## 0.5 0.5

Marquage stochastique des caractères.

Calculons maintenant les fréquences d’état à partir des marquages stochastiques pour chaque noeud interne. Ce sont nos probabilités postérieures pour les noeuds.

Fonction pour calculer les états:

Apliquer la fonction à tous les arbres et tracer l’arbre et les probabilités postérieures aux nœuds.

probabilités postérieures aux nœuds

Maximum de vraisemblance

La commande ace de la librairie ape peut reconstruire des états ancestraux pour des caractères discrets en utilisant le maximum de vraisemblance (Pagel 1994 Detecting correlated evolution on phylogenies: a general method for the comparative analysis of discrete characters. Proceedings of the Royal Society of London. Series B. Biological Sciences, 255, 37–45.).

Par défaut, la probabilité que les différents états ancestraux des caractères discrets sont calculés à l’aide d’une procédure d’estimation conjointe (joint) en utilisant une procédure semblable à celle décrite dans Pupko et al. (2000). Si “marginal = VRAI”, une procédure d’estimation marginale est utilisée. Avec cette méthode, les valeurs de vraisemblance à un nœud donné sont calculées en utilisant uniquement les informations provenant des feuilles (et des branches) descendantes de ce nœud. Avec l’estimation conjointe, toutes les informations sont utilisées pour chaque nœud.

La mise en œuvre actuelle de l’estimation conjointe utilise un algorithme “en deux passes” qui est beaucoup plus rapide que l’approche stochastique alors que les estimations des deux méthodes sont très proches.

Si l’option CI = TRUE est utilisée, alors la probabilité de chaque état ancestral est retournée pour chaque noeud dans une matrice appelée lik.anc.

obj<-anc.Bayes(tree,x,ngen=10000) # sample ancestral states print(obj,printlen=20) ## estimates

Remarque: la fonction fitDiscrete de geiger utilise la même syntaxe que ace et donne des résultats similaires, mais les valeurs de log-vraisemblance sont mises à l’échelle différemment cependant le test du rapport de vraisemblance reste le même.

Exemple motif 3 (“0000001100101111”) avec: * un modèle à un paramètre (ER), * un modèle symétrique (SYM) , * un modèle où tous les taux sont différents (ARD).

## 0111111100000000

Maximum de vraisemblance

La sortie du modèle ER correspond à celle du modèle SYM. Pour un caractère binaire, ces modèles sont identiques, bien qu’ils soient distincts pour les caractères à trois états ou plus. Pour un caractère à trois états, ER est un modèle à un paramètre, SYM un modèle à trois paramètres et ARD un modèle à six paramètres.

Le modèle ARD donne généralement la probabilité la plus élevée. Cependant, il inclut aussi plus de paramètres que le modèle ER. Il est bien connu que l’ajout de paramètres à un modèle augmente généralement sa probabilité. Pour déterminer si l’utilisation du modèle ARD est appropriée, vous devez exécuter un test de vraisemblance.

Test de vraisemblance

La différence de logarithme de probabilité entre deux modèles est distribuée sous forme de chi2, avec des degrés de liberté égaux au nombre de paramètres qui diffèrent entre les modèles. Ainsi, un test de vraisemblance de ces modèles demande si la différence de vraisemblance est suffisamment grande pour se situer dans la queue la plus à droite de la distribution du chi2.

La fonction pchisq(value, df) donne le pourcentage de la fonction de distribution cumulative pour chi2 qui se trouve à gauche de la valeur donnée pour un degré de liberté souhaité(df). Ainsi,

## [1] 0.2827516

Boucle sur tous les motifs

Par curiosité nous allons réaliser le test pour tous les motifs et imprimer les valeurs les plus significatives.

Table: Test de vraisemblance (pvalue < 0.01).
pattern	er loglik	ard loglik	p-value
0	0	0	0
1111111111110110	-9.056	-5.335	0.00638
1111011111111110	-9.859	-5.335	0.00263
1111111110111111	-6.677	-3.048	0.00706
1111101111111101	-9.661	-5.335	0.00327
1111111111011011	-9.434	-5.284	0.00397
1111111101111111	-6.698	-3.048	0.00689
0000000000001001	-9.056	-5.335	0.00638
0000000000010001	-8.622	-5.259	0.00951
0000000000001101	-9.798	-6.461	0.00978
0000000000000101	-9.051	-5.335	0.00641
0010100000000000	-8.757	-5.335	0.00889
1000100000000000	-9.186	-5.335	0.00552
0000000010000000	-6.698	-3.048	0.00689
1000000010000000	-9.268	-5.335	0.00504
0000000000110000	-8.529	-5.198	0.00984
0000000000100100	-9.434	-5.284	0.00397
0000000000101000	-9.437	-5.284	0.00395
0000000000011000	-9.033	-5.187	0.00555
0000000000001010	-8.931	-5.335	0.00732
0000000001010010	-10.397	-7.028	0.00944
0000001001010000	-11.211	-7.028	0.00383
0000000001000000	-6.677	-3.048	0.00706
0010010001000000	-10.397	-7.028	0.00944
1000000001000000	-9.252	-5.335	0.00513
0010000001000000	-8.833	-5.335	0.00817
0000000000010100	-9.031	-5.189	0.00557
1000010000000100	-10.369	-7.028	0.00974
0000010000000100	-10.64	-5.335	0.00113
0100010000000100	-10.369	-7.028	0.00974
0000010000010100	-10.369	-7.028	0.00974
0100010000000000	-9.013	-5.335	0.00668
1000010000000000	-9.112	-5.335	0.00599
0010010000000000	-8.683	-5.335	0.00966

Fonction pour annoter les états des feuilles de l’arbre

Les inférences des états du caractère au nœuds sont calculées sur les noeuds interne de l’arbre, nous ajoutons ici les noeuds feuilles.

Exemple d’appel de la fonction

Concaténation des deux matrices (feuilles et noeuds)

Fonction pour annoter les transitions d’états sur les branches de l’arbre

Une transition d’état est identifiée sur une branche (edge) si l’état du caractère présent au nœud parent est différent de l’état du caractère présent au nœud fils.

En entrée l’arbre tr et les états du caractère inférés aux noeuds (edge_states).
En sortie une liste avec les listes de transitions (type de tansition, direction et une couleur).

Tracé pour chaque motif

## 1 parent 17 1 -> 0 child 18 - 
## 16 parent 24 1 -> 0 child 25 - -

Gain perte de gènes (ER)

## 12 parent 21 0 -> 1 child 23 + 
## 17 parent 25 1 -> 0 child 26 + - 
## 24 parent 29 1 -> 0 child 12 + - -

Gain perte de gènes (ER)

## 15 parent 24 0 -> 1 child 8 +

Gain perte de gènes (ER)

## 4 parent 20 1 -> 0 child 1 - 
## 16 parent 24 1 -> 0 child 25 - -

Gain perte de gènes (ER)

## 20 parent 25 1 -> 0 child 27 - 
## 16 parent 24 1 -> 0 child 25 - 
## 20 parent 25 1 -> 0 child 27 - 
## 27 parent 30 0 -> 1 child 14 - + 
## 30 parent 31 0 -> 1 child 16 - + + 
## 12 parent 21 1 -> 0 child 23 - 
## 15 parent 24 0 -> 1 child 8 - + 
## 15 parent 24 0 -> 1 child 8 + 
## 22 parent 28 0 -> 1 child 11 + + 
## 15 parent 24 0 -> 1 child 8 + 
## 20 parent 25 0 -> 1 child 27 + + 
## 22 parent 28 1 -> 0 child 11 + + - 
## 1 parent 17 0 -> 1 child 18 + 
## 4 parent 20 1 -> 0 child 1 + - 
## 12 parent 21 0 -> 1 child 23 + - + 
## 13 parent 23 1 -> 0 child 7 + - + - 
## 1 parent 17 0 -> 1 child 18 + 
## 4 parent 20 1 -> 0 child 1 + - 
## 15 parent 24 0 -> 1 child 8 + - + 
## 20 parent 25 0 -> 1 child 27 + - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 9 parent 21 1 -> 0 child 22 - 
## 16 parent 24 1 -> 0 child 25 - - 
## 9 parent 21 1 -> 0 child 22 - 
## 6 parent 19 1 -> 0 child 3 - 
## 24 parent 29 1 -> 0 child 12 - - 
## 13 parent 23 1 -> 0 child 7 - 
## 2 parent 18 1 -> 0 child 19 - 
## 5 parent 20 0 -> 1 child 2 - + 
## 30 parent 31 1 -> 0 child 16 - + - 
## 22 parent 28 1 -> 0 child 11 - 
## 6 parent 19 1 -> 0 child 3 - 
## 4 parent 20 1 -> 0 child 1 - 
## 17 parent 25 1 -> 0 child 26 - 
## 10 parent 22 0 -> 1 child 5 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 20 parent 25 1 -> 0 child 27 + + - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 30 parent 31 1 -> 0 child 16 - 
## 26 parent 30 1 -> 0 child 13 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 10 parent 22 1 -> 0 child 5 - 
## 5 parent 20 1 -> 0 child 2 - 
## 9 parent 21 1 -> 0 child 22 - 
## 17 parent 25 1 -> 0 child 26 - - 
## 11 parent 22 1 -> 0 child 6 - 
## 3 parent 19 1 -> 0 child 20 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 15 parent 24 0 -> 1 child 8 - - + 
## 17 parent 25 1 -> 0 child 26 - 
## 29 parent 31 1 -> 0 child 15 - - 
## 1 parent 17 1 -> 0 child 18 - 
## 2 parent 18 1 -> 0 child 19 - 
## 6 parent 19 1 -> 0 child 3 - 
## 7 parent 18 1 -> 0 child 4 - - 
## 10 parent 22 1 -> 0 child 5 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 19 parent 26 1 -> 0 child 10 - 
## 3 parent 19 1 -> 0 child 20 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 5 parent 20 1 -> 0 child 2 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 7 parent 18 0 -> 1 child 4 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 29 parent 31 1 -> 0 child 15 - 
## 5 parent 20 0 -> 1 child 2 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 17 parent 25 0 -> 1 child 26 + + - + 
## 19 parent 26 1 -> 0 child 10 + + - + - 
## 1 parent 17 1 -> 0 child 18 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 24 parent 29 0 -> 1 child 12 - - + 
## 29 parent 31 0 -> 1 child 15 - - + + 
## 26 parent 30 1 -> 0 child 13 - 
## 5 parent 20 0 -> 1 child 2 + 
## 6 parent 19 0 -> 1 child 3 + + 
## 12 parent 21 0 -> 1 child 23 + + + 
## 6 parent 19 1 -> 0 child 3 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 12 parent 21 0 -> 1 child 23 + 
## 17 parent 25 1 -> 0 child 26 + - 
## 20 parent 25 1 -> 0 child 27 - 
## 24 parent 29 0 -> 1 child 12 - + 
## 26 parent 30 0 -> 1 child 13 - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 11 parent 22 1 -> 0 child 6 - 
## 29 parent 31 1 -> 0 child 15 - - 
## 13 parent 23 1 -> 0 child 7 - 
## 17 parent 25 1 -> 0 child 26 - - 
## 7 parent 18 1 -> 0 child 4 - 
## 17 parent 25 1 -> 0 child 26 - 
## 21 parent 27 1 -> 0 child 28 - - 
## 1 parent 17 1 -> 0 child 18 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 15 parent 24 0 -> 1 child 8 - - + 
## 20 parent 25 0 -> 1 child 27 - - + + 
## 24 parent 29 1 -> 0 child 12 - - + + - 
## 20 parent 25 1 -> 0 child 27 - 
## 22 parent 28 0 -> 1 child 11 - + 
## 27 parent 30 0 -> 1 child 14 - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 20 parent 25 0 -> 1 child 27 + - + 
## 23 parent 28 1 -> 0 child 29 + - + - 
## 24 parent 29 0 -> 1 child 12 + - + - + 
## 28 parent 27 1 -> 0 child 31 + - + - + - 
## 29 parent 31 0 -> 1 child 15 + - + - + - + 
## 4 parent 20 0 -> 1 child 1 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 1 parent 17 1 -> 0 child 18 - 
## 6 parent 19 0 -> 1 child 3 - + 
## 15 parent 24 0 -> 1 child 8 + 
## 20 parent 25 0 -> 1 child 27 + + 
## 2 parent 18 1 -> 0 child 19 - 
## 4 parent 20 0 -> 1 child 1 - + 
## 8 parent 17 1 -> 0 child 21 - + - 
## 11 parent 22 0 -> 1 child 6 - + - + 
## 14 parent 23 0 -> 1 child 24 - + - + + 
## 4 parent 20 0 -> 1 child 1 + 
## 15 parent 24 0 -> 1 child 8 + + 
## 24 parent 29 0 -> 1 child 12 + + + 
## 11 parent 22 0 -> 1 child 6 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 2 parent 18 0 -> 1 child 19 + 
## 15 parent 24 0 -> 1 child 8 + + 
## 22 parent 28 0 -> 1 child 11 + + + 
## 14 parent 23 0 -> 1 child 24 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 23 parent 28 0 -> 1 child 29 + - + 
## 26 parent 30 1 -> 0 child 13 + - + - 
## 1 parent 17 1 -> 0 child 18 - 
## 7 parent 18 0 -> 1 child 4 - + 
## 12 parent 21 1 -> 0 child 23 - + - 
## 14 parent 23 0 -> 1 child 24 - + - + 
## 16 parent 24 1 -> 0 child 25 - + - + - 
## 20 parent 25 0 -> 1 child 27 - + - + - + 
## 27 parent 30 1 -> 0 child 14 - + - + - + - 
## 14 parent 23 0 -> 1 child 24 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 30 parent 31 0 -> 1 child 16 + - + 
## 4 parent 20 1 -> 0 child 1 - 
## 5 parent 20 1 -> 0 child 2 - - 
## 6 parent 19 1 -> 0 child 3 - - - 
## 7 parent 18 1 -> 0 child 4 - - - - 
## 9 parent 21 1 -> 0 child 22 - - - - - 
## 17 parent 25 1 -> 0 child 26 - - - - - - 
## 19 parent 26 0 -> 1 child 10 - - - - - - + 
## 23 parent 28 1 -> 0 child 29 - - - - - - + - 
## 29 parent 31 1 -> 0 child 15 - - - - - - + - - 
## 30 parent 31 1 -> 0 child 16 - - - - - - + - - - 
## 4 parent 20 1 -> 0 child 1 - 
## 5 parent 20 1 -> 0 child 2 - - 
## 6 parent 19 1 -> 0 child 3 - - - 
## 7 parent 18 1 -> 0 child 4 - - - - 
## 13 parent 23 1 -> 0 child 7 - - - - - 
## 19 parent 26 1 -> 0 child 10 - - - - - - 
## 24 parent 29 1 -> 0 child 12 - - - - - - - 
## 25 parent 29 1 -> 0 child 30 - - - - - - - - 
## 30 parent 31 1 -> 0 child 16 - - - - - - - - - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 23 parent 28 0 -> 1 child 29 + - + 
## 27 parent 30 1 -> 0 child 14 + - + - 
## 29 parent 31 0 -> 1 child 15 + - + - + 
## 14 parent 23 0 -> 1 child 24 + 
## 1 parent 17 1 -> 0 child 18 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 15 parent 24 0 -> 1 child 8 + 
## 16 parent 24 0 -> 1 child 25 + + 
## 20 parent 25 1 -> 0 child 27 + + - 
## 12 parent 21 1 -> 0 child 23 - 
## 14 parent 23 0 -> 1 child 24 - + 
## 20 parent 25 1 -> 0 child 27 - + - 
## 23 parent 28 0 -> 1 child 29 - + - + 
## 24 parent 29 1 -> 0 child 12 - + - + - 
## 29 parent 31 0 -> 1 child 15 - + - + - + 
## 12 parent 21 1 -> 0 child 23 - 
## 14 parent 23 0 -> 1 child 24 - + 
## 16 parent 24 1 -> 0 child 25 - + - 
## 23 parent 28 0 -> 1 child 29 - + - + 
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## 9 parent 21 1 -> 0 child 22 - - - - 
## 11 parent 22 0 -> 1 child 6 - - - - + 
## 13 parent 23 1 -> 0 child 7 - - - - + - 
## 15 parent 24 1 -> 0 child 8 - - - - + - - 
## 17 parent 25 1 -> 0 child 26 - - - - + - - - 
## 19 parent 26 0 -> 1 child 10 - - - - + - - - + 
## 22 parent 28 1 -> 0 child 11 - - - - + - - - + - 
## 24 parent 29 1 -> 0 child 12 - - - - + - - - + - - 
## 25 parent 29 1 -> 0 child 30 - - - - + - - - + - - - 
## 29 parent 31 1 -> 0 child 15 - - - - + - - - + - - - - 
## 30 parent 31 1 -> 0 child 16 - - - - + - - - + - - - - - 
## 7 parent 18 0 -> 1 child 4 + 
## 19 parent 26 0 -> 1 child 10 + + 
## 4 parent 20 0 -> 1 child 1 + 
## 19 parent 26 0 -> 1 child 10 + + 
## 6 parent 19 0 -> 1 child 3 + 
## 7 parent 18 0 -> 1 child 4 + + 
## 19 parent 26 0 -> 1 child 10 + + + 
## 6 parent 19 0 -> 1 child 3 + 
## 19 parent 26 0 -> 1 child 10 + + 
## 4 parent 20 1 -> 0 child 1 - 
## 7 parent 18 1 -> 0 child 4 - - 
## 9 parent 21 1 -> 0 child 22 - - - 
## 11 parent 22 0 -> 1 child 6 - - - + 
## 13 parent 23 1 -> 0 child 7 - - - + - 
## 14 parent 23 1 -> 0 child 24 - - - + - - 
## 16 parent 24 0 -> 1 child 25 - - - + - - + 
## 17 parent 25 1 -> 0 child 26 - - - + - - + - 
## 19 parent 26 0 -> 1 child 10 - - - + - - + - + 
## 22 parent 28 1 -> 0 child 11 - - - + - - + - + - 
## 23 parent 28 1 -> 0 child 29 - - - + - - + - + - - 
## 29 parent 31 1 -> 0 child 15 - - - + - - + - + - - - 
## 30 parent 31 1 -> 0 child 16 - - - + - - + - + - - - - 
## 27 parent 30 0 -> 1 child 14 + 
## 24 parent 29 0 -> 1 child 12 + 
## 27 parent 30 0 -> 1 child 14 + + 
## 2 parent 18 0 -> 1 child 19 + 
## 5 parent 20 1 -> 0 child 2 + - 
## 6 parent 19 1 -> 0 child 3 + - - 
## 11 parent 22 0 -> 1 child 6 + - - + 
## 27 parent 30 0 -> 1 child 14 + - - + + 
## 11 parent 22 0 -> 1 child 6 + 
## 27 parent 30 0 -> 1 child 14 + + 
## 3 parent 19 0 -> 1 child 20 + 
## 27 parent 30 0 -> 1 child 14 + + 
## 5 parent 20 0 -> 1 child 2 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 27 parent 30 0 -> 1 child 14 + + + 
## 11 parent 22 0 -> 1 child 6 + 
## 23 parent 28 0 -> 1 child 29 + + 
## 26 parent 30 1 -> 0 child 13 + + - 
## 2 parent 18 0 -> 1 child 19 + 
## 23 parent 28 0 -> 1 child 29 + + 
## 26 parent 30 1 -> 0 child 13 + + - 
## 24 parent 29 0 -> 1 child 12 + 
## 1 parent 17 0 -> 1 child 18 + 
## 24 parent 29 0 -> 1 child 12 + + 
## 11 parent 22 0 -> 1 child 6 + 
## 5 parent 20 0 -> 1 child 2 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 3 parent 19 0 -> 1 child 20 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 10 parent 22 1 -> 0 child 5 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 4 parent 20 0 -> 1 child 1 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 1 parent 17 0 -> 1 child 18 + 
## 2 parent 18 1 -> 0 child 19 + - 
## 6 parent 19 0 -> 1 child 3 + - + 
## 11 parent 22 0 -> 1 child 6 + - + + 
## 1 parent 17 0 -> 1 child 18 + 
## 2 parent 18 1 -> 0 child 19 + - 
## 5 parent 20 0 -> 1 child 2 + - + 
## 11 parent 22 0 -> 1 child 6 + - + + 
## 1 parent 17 0 -> 1 child 18 + 
## 7 parent 18 1 -> 0 child 4 + - 
## 11 parent 22 0 -> 1 child 6 + - + 
## 6 parent 19 0 -> 1 child 3 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 7 parent 18 0 -> 1 child 4 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 2 parent 18 0 -> 1 child 19 + 
## 5 parent 20 1 -> 0 child 2 + - 
## 11 parent 22 0 -> 1 child 6 + - + 
## 1 parent 17 0 -> 1 child 18 + 
## 2 parent 18 1 -> 0 child 19 + - 
## 4 parent 20 0 -> 1 child 1 + - + 
## 11 parent 22 0 -> 1 child 6 + - + + 
## 4 parent 20 0 -> 1 child 1 + 
## 6 parent 19 0 -> 1 child 3 + + 
## 6 parent 19 0 -> 1 child 3 + 
## 5 parent 20 0 -> 1 child 2 + 
## 6 parent 19 0 -> 1 child 3 + + 
## 1 parent 17 0 -> 1 child 18 + 
## 2 parent 18 0 -> 1 child 19 + 
## 6 parent 19 0 -> 1 child 3 + 
## 7 parent 18 0 -> 1 child 4 + + 
## 1 parent 17 0 -> 1 child 18 + 
## 4 parent 20 1 -> 0 child 1 + - 
## 1 parent 17 0 -> 1 child 18 + 
## 5 parent 20 1 -> 0 child 2 + - 
## 4 parent 20 0 -> 1 child 1 + 
## 3 parent 19 0 -> 1 child 20 + 
## 4 parent 20 0 -> 1 child 1 + 
## 7 parent 18 0 -> 1 child 4 + + 
## 1 parent 17 0 -> 1 child 18 + 
## 6 parent 19 1 -> 0 child 3 + - 
## 7 parent 18 0 -> 1 child 4 + 
## 5 parent 20 0 -> 1 child 2 + 
## 7 parent 18 0 -> 1 child 4 + +

Flux de gènes

## 1 parent 17 1 -> 0 child 18 - 
## 16 parent 24 1 -> 0 child 25 - - 
## 12 parent 21 0 -> 1 child 23 + 
## 17 parent 25 1 -> 0 child 26 + - 
## 24 parent 29 1 -> 0 child 12 + - - 
## 15 parent 24 0 -> 1 child 8 + 
## 4 parent 20 1 -> 0 child 1 - 
## 16 parent 24 1 -> 0 child 25 - - 
## 20 parent 25 1 -> 0 child 27 - 
## 16 parent 24 1 -> 0 child 25 - 
## 20 parent 25 1 -> 0 child 27 - 
## 27 parent 30 0 -> 1 child 14 - + 
## 30 parent 31 0 -> 1 child 16 - + + 
## 12 parent 21 1 -> 0 child 23 - 
## 15 parent 24 0 -> 1 child 8 - + 
## 15 parent 24 0 -> 1 child 8 + 
## 22 parent 28 0 -> 1 child 11 + + 
## 15 parent 24 0 -> 1 child 8 + 
## 20 parent 25 0 -> 1 child 27 + + 
## 22 parent 28 1 -> 0 child 11 + + - 
## 1 parent 17 0 -> 1 child 18 + 
## 4 parent 20 1 -> 0 child 1 + - 
## 12 parent 21 0 -> 1 child 23 + - + 
## 13 parent 23 1 -> 0 child 7 + - + - 
## 1 parent 17 0 -> 1 child 18 + 
## 4 parent 20 1 -> 0 child 1 + - 
## 15 parent 24 0 -> 1 child 8 + - + 
## 20 parent 25 0 -> 1 child 27 + - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 9 parent 21 1 -> 0 child 22 - 
## 16 parent 24 1 -> 0 child 25 - - 
## 9 parent 21 1 -> 0 child 22 - 
## 6 parent 19 1 -> 0 child 3 - 
## 24 parent 29 1 -> 0 child 12 - - 
## 13 parent 23 1 -> 0 child 7 - 
## 2 parent 18 1 -> 0 child 19 - 
## 5 parent 20 0 -> 1 child 2 - + 
## 30 parent 31 1 -> 0 child 16 - + - 
## 22 parent 28 1 -> 0 child 11 - 
## 6 parent 19 1 -> 0 child 3 - 
## 4 parent 20 1 -> 0 child 1 - 
## 17 parent 25 1 -> 0 child 26 - 
## 10 parent 22 0 -> 1 child 5 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 20 parent 25 1 -> 0 child 27 + + - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 30 parent 31 1 -> 0 child 16 - 
## 26 parent 30 1 -> 0 child 13 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 10 parent 22 1 -> 0 child 5 - 
## 5 parent 20 1 -> 0 child 2 - 
## 9 parent 21 1 -> 0 child 22 - 
## 17 parent 25 1 -> 0 child 26 - - 
## 11 parent 22 1 -> 0 child 6 - 
## 3 parent 19 1 -> 0 child 20 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 15 parent 24 0 -> 1 child 8 - - + 
## 17 parent 25 1 -> 0 child 26 - 
## 29 parent 31 1 -> 0 child 15 - - 
## 1 parent 17 1 -> 0 child 18 - 
## 2 parent 18 1 -> 0 child 19 - 
## 6 parent 19 1 -> 0 child 3 - 
## 7 parent 18 1 -> 0 child 4 - - 
## 10 parent 22 1 -> 0 child 5 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 19 parent 26 1 -> 0 child 10 - 
## 3 parent 19 1 -> 0 child 20 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 5 parent 20 1 -> 0 child 2 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 7 parent 18 0 -> 1 child 4 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 29 parent 31 1 -> 0 child 15 - 
## 5 parent 20 0 -> 1 child 2 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 17 parent 25 0 -> 1 child 26 + + - + 
## 19 parent 26 1 -> 0 child 10 + + - + - 
## 1 parent 17 1 -> 0 child 18 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 24 parent 29 0 -> 1 child 12 - - + 
## 29 parent 31 0 -> 1 child 15 - - + + 
## 26 parent 30 1 -> 0 child 13 - 
## 5 parent 20 0 -> 1 child 2 + 
## 6 parent 19 0 -> 1 child 3 + + 
## 12 parent 21 0 -> 1 child 23 + + + 
## 6 parent 19 1 -> 0 child 3 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 12 parent 21 0 -> 1 child 23 + 
## 17 parent 25 1 -> 0 child 26 + - 
## 20 parent 25 1 -> 0 child 27 - 
## 24 parent 29 0 -> 1 child 12 - + 
## 26 parent 30 0 -> 1 child 13 - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 11 parent 22 1 -> 0 child 6 - 
## 29 parent 31 1 -> 0 child 15 - - 
## 13 parent 23 1 -> 0 child 7 - 
## 17 parent 25 1 -> 0 child 26 - - 
## 7 parent 18 1 -> 0 child 4 - 
## 17 parent 25 1 -> 0 child 26 - 
## 21 parent 27 1 -> 0 child 28 - - 
## 1 parent 17 1 -> 0 child 18 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 15 parent 24 0 -> 1 child 8 - - + 
## 20 parent 25 0 -> 1 child 27 - - + + 
## 24 parent 29 1 -> 0 child 12 - - + + - 
## 20 parent 25 1 -> 0 child 27 - 
## 22 parent 28 0 -> 1 child 11 - + 
## 27 parent 30 0 -> 1 child 14 - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 20 parent 25 0 -> 1 child 27 + - + 
## 23 parent 28 1 -> 0 child 29 + - + - 
## 24 parent 29 0 -> 1 child 12 + - + - + 
## 28 parent 27 1 -> 0 child 31 + - + - + - 
## 29 parent 31 0 -> 1 child 15 + - + - + - + 
## 4 parent 20 0 -> 1 child 1 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 1 parent 17 1 -> 0 child 18 - 
## 6 parent 19 0 -> 1 child 3 - + 
## 15 parent 24 0 -> 1 child 8 + 
## 20 parent 25 0 -> 1 child 27 + + 
## 2 parent 18 1 -> 0 child 19 - 
## 4 parent 20 0 -> 1 child 1 - + 
## 8 parent 17 1 -> 0 child 21 - + - 
## 11 parent 22 0 -> 1 child 6 - + - + 
## 14 parent 23 0 -> 1 child 24 - + - + + 
## 4 parent 20 0 -> 1 child 1 + 
## 15 parent 24 0 -> 1 child 8 + + 
## 24 parent 29 0 -> 1 child 12 + + + 
## 11 parent 22 0 -> 1 child 6 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 2 parent 18 0 -> 1 child 19 + 
## 15 parent 24 0 -> 1 child 8 + + 
## 22 parent 28 0 -> 1 child 11 + + + 
## 14 parent 23 0 -> 1 child 24 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 23 parent 28 0 -> 1 child 29 + - + 
## 26 parent 30 1 -> 0 child 13 + - + - 
## 1 parent 17 1 -> 0 child 18 - 
## 7 parent 18 0 -> 1 child 4 - + 
## 12 parent 21 1 -> 0 child 23 - + - 
## 14 parent 23 0 -> 1 child 24 - + - + 
## 16 parent 24 1 -> 0 child 25 - + - + - 
## 20 parent 25 0 -> 1 child 27 - + - + - + 
## 27 parent 30 1 -> 0 child 14 - + - + - + - 
## 14 parent 23 0 -> 1 child 24 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 30 parent 31 0 -> 1 child 16 + - + 
## 4 parent 20 1 -> 0 child 1 - 
## 5 parent 20 1 -> 0 child 2 - - 
## 6 parent 19 1 -> 0 child 3 - - - 
## 7 parent 18 1 -> 0 child 4 - - - - 
## 9 parent 21 1 -> 0 child 22 - - - - - 
## 17 parent 25 1 -> 0 child 26 - - - - - - 
## 19 parent 26 0 -> 1 child 10 - - - - - - + 
## 23 parent 28 1 -> 0 child 29 - - - - - - + - 
## 29 parent 31 1 -> 0 child 15 - - - - - - + - - 
## 30 parent 31 1 -> 0 child 16 - - - - - - + - - - 
## 4 parent 20 1 -> 0 child 1 - 
## 5 parent 20 1 -> 0 child 2 - - 
## 6 parent 19 1 -> 0 child 3 - - - 
## 7 parent 18 1 -> 0 child 4 - - - - 
## 13 parent 23 1 -> 0 child 7 - - - - - 
## 19 parent 26 1 -> 0 child 10 - - - - - - 
## 24 parent 29 1 -> 0 child 12 - - - - - - - 
## 25 parent 29 1 -> 0 child 30 - - - - - - - - 
## 30 parent 31 1 -> 0 child 16 - - - - - - - - - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 23 parent 28 0 -> 1 child 29 + - + 
## 27 parent 30 1 -> 0 child 14 + - + - 
## 29 parent 31 0 -> 1 child 15 + - + - + 
## 14 parent 23 0 -> 1 child 24 + 
## 1 parent 17 1 -> 0 child 18 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 15 parent 24 0 -> 1 child 8 + 
## 16 parent 24 0 -> 1 child 25 + + 
## 20 parent 25 1 -> 0 child 27 + + - 
## 12 parent 21 1 -> 0 child 23 - 
## 14 parent 23 0 -> 1 child 24 - + 
## 20 parent 25 1 -> 0 child 27 - + - 
## 23 parent 28 0 -> 1 child 29 - + - + 
## 24 parent 29 1 -> 0 child 12 - + - + - 
## 29 parent 31 0 -> 1 child 15 - + - + - + 
## 12 parent 21 1 -> 0 child 23 - 
## 14 parent 23 0 -> 1 child 24 - + 
## 16 parent 24 1 -> 0 child 25 - + - 
## 23 parent 28 0 -> 1 child 29 - + - + 
## 27 parent 30 1 -> 0 child 14 - + - + - 
## 28 parent 27 0 -> 1 child 31 - + - + - + 
## 29 parent 31 1 -> 0 child 15 - + - + - + - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 22 parent 28 0 -> 1 child 11 + - + 
## 21 parent 27 1 -> 0 child 28 - 
## 22 parent 28 0 -> 1 child 11 - + 
## 30 parent 31 1 -> 0 child 16 - + - 
## 4 parent 20 1 -> 0 child 1 - 
## 6 parent 19 1 -> 0 child 3 - - 
## 9 parent 21 1 -> 0 child 22 - - - 
## 13 parent 23 1 -> 0 child 7 - - - - 
## 17 parent 25 1 -> 0 child 26 - - - - - 
## 19 parent 26 0 -> 1 child 10 - - - - - + 
## 24 parent 29 1 -> 0 child 12 - - - - - + - 
## 30 parent 31 1 -> 0 child 16 - - - - - + - - 
## 4 parent 20 1 -> 0 child 1 - 
## 6 parent 19 1 -> 0 child 3 - - 
## 7 parent 18 1 -> 0 child 4 - - - 
## 9 parent 21 1 -> 0 child 22 - - - - 
## 13 parent 23 1 -> 0 child 7 - - - - - 
## 17 parent 25 1 -> 0 child 26 - - - - - - 
## 19 parent 26 0 -> 1 child 10 - - - - - - + 
## 22 parent 28 1 -> 0 child 11 - - - - - - + - 
## 24 parent 29 1 -> 0 child 12 - - - - - - + - - 
## 27 parent 30 1 -> 0 child 14 - - - - - - + - - - 
## 30 parent 31 1 -> 0 child 16 - - - - - - + - - - - 
## 12 parent 21 0 -> 1 child 23 + 
## 25 parent 29 1 -> 0 child 30 + - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 24 parent 29 0 -> 1 child 12 + - + 
## 14 parent 23 0 -> 1 child 24 + 
## 21 parent 27 1 -> 0 child 28 + - 
## 25 parent 29 0 -> 1 child 30 + - + 
## 7 parent 18 1 -> 0 child 4 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 15 parent 24 0 -> 1 child 8 - - + 
## 5 parent 20 1 -> 0 child 2 - 
## 7 parent 18 1 -> 0 child 4 - - 
## 2 parent 18 1 -> 0 child 19 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 15 parent 24 0 -> 1 child 8 - - + 
## 14 parent 23 0 -> 1 child 24 + 
## 19 parent 26 1 -> 0 child 10 + - 
## 1 parent 17 1 -> 0 child 18 - 
## 13 parent 23 1 -> 0 child 7 - - 
## 2 parent 18 0 -> 1 child 19 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 9 parent 21 1 -> 0 child 22 - 
## 29 parent 31 1 -> 0 child 15 - - 
## 12 parent 21 0 -> 1 child 23 + 
## 22 parent 28 1 -> 0 child 11 + - 
## 29 parent 31 1 -> 0 child 15 + - - 
## 24 parent 29 1 -> 0 child 12 - 
## 7 parent 18 0 -> 1 child 4 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 22 parent 28 1 -> 0 child 11 - 
## 27 parent 30 1 -> 0 child 14 - - 
## 3 parent 19 1 -> 0 child 20 - 
## 5 parent 20 0 -> 1 child 2 - + 
## 23 parent 28 1 -> 0 child 29 - + - 
## 24 parent 29 0 -> 1 child 12 - + - + 
## 29 parent 31 1 -> 0 child 15 - + - + - 
## 15 parent 24 0 -> 1 child 8 + 
## 21 parent 27 0 -> 1 child 28 + + 
## 2 parent 18 0 -> 1 child 19 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 17 parent 25 1 -> 0 child 26 + + - 
## 17 parent 25 1 -> 0 child 26 - 
## 24 parent 29 1 -> 0 child 12 - - 
## 1 parent 17 0 -> 1 child 18 + 
## 2 parent 18 1 -> 0 child 19 + - 
## 6 parent 19 0 -> 1 child 3 + - + 
## 11 parent 22 0 -> 1 child 6 + - + + 
## 14 parent 23 0 -> 1 child 24 + - + + + 
## 20 parent 25 1 -> 0 child 27 + - + + + - 
## 23 parent 28 0 -> 1 child 29 + - + + + - + 
## 27 parent 30 1 -> 0 child 14 + - + + + - + - 
## 29 parent 31 0 -> 1 child 15 + - + + + - + - + 
## 6 parent 19 1 -> 0 child 3 - 
## 13 parent 23 1 -> 0 child 7 - - 
## 20 parent 25 1 -> 0 child 27 - - - 
## 5 parent 20 1 -> 0 child 2 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 5 parent 20 0 -> 1 child 2 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 30 parent 31 1 -> 0 child 16 + + - 
## 7 parent 18 1 -> 0 child 4 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 7 parent 18 0 -> 1 child 4 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 29 parent 31 1 -> 0 child 15 + + - 
## 23 parent 28 1 -> 0 child 29 - 
## 24 parent 29 1 -> 0 child 12 - 
## 29 parent 31 1 -> 0 child 15 - - 
## 15 parent 24 0 -> 1 child 8 + 
## 19 parent 26 0 -> 1 child 10 + + 
## 20 parent 25 1 -> 0 child 27 - 
## 23 parent 28 0 -> 1 child 29 - + 
## 26 parent 30 1 -> 0 child 13 - + - 
## 28 parent 27 0 -> 1 child 31 - + - + 
## 29 parent 31 1 -> 0 child 15 - + - + - 
## 7 parent 18 1 -> 0 child 4 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 14 parent 23 0 -> 1 child 24 - - + 
## 27 parent 30 1 -> 0 child 14 - - + - 
## 28 parent 27 1 -> 0 child 31 - - + - - 
## 29 parent 31 0 -> 1 child 15 - - + - - + 
## 9 parent 21 1 -> 0 child 22 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 9 parent 21 1 -> 0 child 22 - 
## 13 parent 23 1 -> 0 child 7 - - 
## 5 parent 20 1 -> 0 child 2 - 
## 9 parent 21 1 -> 0 child 22 - - 
## 11 parent 22 0 -> 1 child 6 - - + 
## 12 parent 21 1 -> 0 child 23 - - + - 
## 14 parent 23 0 -> 1 child 24 - - + - + 
## 20 parent 25 1 -> 0 child 27 - - + - + - 
## 4 parent 20 1 -> 0 child 1 - 
## 6 parent 19 1 -> 0 child 3 - - 
## 9 parent 21 1 -> 0 child 22 - - - 
## 13 parent 23 1 -> 0 child 7 - - - - 
## 17 parent 25 1 -> 0 child 26 - - - - - 
## 19 parent 26 0 -> 1 child 10 - - - - - + 
## 24 parent 29 1 -> 0 child 12 - - - - - + - 
## 25 parent 29 1 -> 0 child 30 - - - - - + - - 
## 27 parent 30 0 -> 1 child 14 - - - - - + - - + 
## 12 parent 21 0 -> 1 child 23 + 
## 27 parent 30 1 -> 0 child 14 + - 
## 30 parent 31 1 -> 0 child 16 + - - 
## 12 parent 21 0 -> 1 child 23 + 
## 17 parent 25 1 -> 0 child 26 + - 
## 29 parent 31 1 -> 0 child 15 + - - 
## 4 parent 20 1 -> 0 child 1 - 
## 7 parent 18 1 -> 0 child 4 - - 
## 2 parent 18 0 -> 1 child 19 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 20 parent 25 1 -> 0 child 27 + + - 
## 1 parent 17 1 -> 0 child 18 - 
## 27 parent 30 1 -> 0 child 14 - - 
## 20 parent 25 1 -> 0 child 27 - 
## 22 parent 28 0 -> 1 child 11 - + 
## 9 parent 21 0 -> 1 child 22 + 
## 15 parent 24 0 -> 1 child 8 + + 
## 9 parent 21 1 -> 0 child 22 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 5 parent 20 1 -> 0 child 2 - 
## 7 parent 18 1 -> 0 child 4 - - 
## 29 parent 31 1 -> 0 child 15 - - - 
## 4 parent 20 1 -> 0 child 1 - 
## 12 parent 21 1 -> 0 child 23 - - 
## 14 parent 23 0 -> 1 child 24 - - + 
## 16 parent 24 1 -> 0 child 25 - - + - 
## 20 parent 25 0 -> 1 child 27 - - + - + 
## 15 parent 24 0 -> 1 child 8 + 
## 28 parent 27 0 -> 1 child 31 + + 
## 14 parent 23 0 -> 1 child 24 + 
## 28 parent 27 1 -> 0 child 31 + - 
## 14 parent 23 0 -> 1 child 24 + 
## 22 parent 28 1 -> 0 child 11 + - 
## 6 parent 19 0 -> 1 child 3 + 
## 7 parent 18 0 -> 1 child 4 + + 
## 12 parent 21 0 -> 1 child 23 + + + 
## 3 parent 19 0 -> 1 child 20 + 
## 5 parent 20 1 -> 0 child 2 + - 
## 14 parent 23 0 -> 1 child 24 + - + 
## 16 parent 24 1 -> 0 child 25 + - + - 
## 27 parent 30 0 -> 1 child 14 + - + - + 
## 28 parent 27 0 -> 1 child 31 + - + - + + 
## 29 parent 31 1 -> 0 child 15 + - + - + + - 
## 17 parent 25 1 -> 0 child 26 - 
## 24 parent 29 1 -> 0 child 12 - - 
## 29 parent 31 1 -> 0 child 15 - - - 
## 15 parent 24 0 -> 1 child 8 + 
## 24 parent 29 0 -> 1 child 12 + + 
## 3 parent 19 1 -> 0 child 20 - 
## 30 parent 31 1 -> 0 child 16 - - 
## 14 parent 23 0 -> 1 child 24 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 26 parent 30 0 -> 1 child 13 + - + 
## 28 parent 27 0 -> 1 child 31 + - + + 
## 29 parent 31 1 -> 0 child 15 + - + + - 
## 15 parent 24 0 -> 1 child 8 + 
## 26 parent 30 0 -> 1 child 13 + + 
## 12 parent 21 0 -> 1 child 23 + 
## 24 parent 29 1 -> 0 child 12 + - 
## 30 parent 31 1 -> 0 child 16 + - - 
## 15 parent 24 0 -> 1 child 8 + 
## 21 parent 27 0 -> 1 child 28 + + 
## 25 parent 29 1 -> 0 child 30 + + - 
## 12 parent 21 1 -> 0 child 23 - 
## 14 parent 23 0 -> 1 child 24 - + 
## 21 parent 27 1 -> 0 child 28 - + - 
## 23 parent 28 0 -> 1 child 29 - + - + 
## 26 parent 30 1 -> 0 child 13 - + - + - 
## 5 parent 20 0 -> 1 child 2 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 20 parent 25 0 -> 1 child 27 + + - + 
## 23 parent 28 1 -> 0 child 29 + + - + - 
## 24 parent 29 0 -> 1 child 12 + + - + - + 
## 29 parent 31 1 -> 0 child 15 + + - + - + - 
## 15 parent 24 0 -> 1 child 8 + 
## 30 parent 31 0 -> 1 child 16 + + 
## 11 parent 22 0 -> 1 child 6 + 
## 14 parent 23 0 -> 1 child 24 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 26 parent 30 0 -> 1 child 13 + + - + 
## 28 parent 27 0 -> 1 child 31 + + - + + 
## 29 parent 31 1 -> 0 child 15 + + - + + - 
## 15 parent 24 0 -> 1 child 8 + 
## 25 parent 29 0 -> 1 child 30 + + 
## 14 parent 23 0 -> 1 child 24 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 20 parent 25 0 -> 1 child 27 + - + 
## 21 parent 27 1 -> 0 child 28 + - + - 
## 23 parent 28 0 -> 1 child 29 + - + - + 
## 27 parent 30 1 -> 0 child 14 + - + - + - 
## 5 parent 20 1 -> 0 child 2 - 
## 6 parent 19 1 -> 0 child 3 - - 
## 20 parent 25 1 -> 0 child 27 - 
## 24 parent 29 0 -> 1 child 12 - + 
## 5 parent 20 0 -> 1 child 2 + 
## 7 parent 18 0 -> 1 child 4 + + 
## 12 parent 21 0 -> 1 child 23 + + + 
## 17 parent 25 1 -> 0 child 26 + + + - 
## 1 parent 17 1 -> 0 child 18 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 22 parent 28 0 -> 1 child 11 - - + 
## 2 parent 18 1 -> 0 child 19 - 
## 4 parent 20 0 -> 1 child 1 - + 
## 22 parent 28 1 -> 0 child 11 - + - 
## 12 parent 21 0 -> 1 child 23 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 22 parent 28 0 -> 1 child 11 + - + 
## 10 parent 22 0 -> 1 child 5 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 4 parent 20 0 -> 1 child 1 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 16 parent 24 1 -> 0 child 25 + + - 
## 14 parent 23 0 -> 1 child 24 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 26 parent 30 0 -> 1 child 13 + - + 
## 29 parent 31 0 -> 1 child 15 + - + + 
## 5 parent 20 0 -> 1 child 2 + 
## 15 parent 24 0 -> 1 child 8 + + 
## 29 parent 31 0 -> 1 child 15 + + + 
## 1 parent 17 0 -> 1 child 18 + 
## 12 parent 21 0 -> 1 child 23 + + 
## 20 parent 25 1 -> 0 child 27 + + - 
## 21 parent 27 0 -> 1 child 28 + + - + 
## 24 parent 29 1 -> 0 child 12 + + - + - 
## 1 parent 17 1 -> 0 child 18 - 
## 28 parent 27 1 -> 0 child 31 - - 
## 1 parent 17 1 -> 0 child 18 - 
## 25 parent 29 1 -> 0 child 30 - - 
## 15 parent 24 0 -> 1 child 8 + 
## 20 parent 25 0 -> 1 child 27 + + 
## 25 parent 29 1 -> 0 child 30 + + - 
## 12 parent 21 0 -> 1 child 23 + 
## 20 parent 25 1 -> 0 child 27 + - 
## 24 parent 29 0 -> 1 child 12 + - + 
## 29 parent 31 0 -> 1 child 15 + - + + 
## 12 parent 21 0 -> 1 child 23 + 
## 28 parent 27 1 -> 0 child 31 + - 
## 7 parent 18 0 -> 1 child 4 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 15 parent 24 0 -> 1 child 8 + + + 
## 7 parent 18 1 -> 0 child 4 - 
## 20 parent 25 1 -> 0 child 27 - - 
## 22 parent 28 0 -> 1 child 11 - - + 
## 5 parent 20 0 -> 1 child 2 + 
## 11 parent 22 0 -> 1 child 6 + + 
## 12 parent 21 0 -> 1 child 23 + + + 
## 15 parent 24 0 -> 1 child 8 + 
## 21 parent 27 0 -> 1 child 28 + + 
## 24 parent 29 1 -> 0 child 12 + + - 
## 1 parent 17 0 -> 1 child 18 + 
## 4 parent 20 1 -> 0 child 1 + - 
## 12 parent 21 0 -> 1 child 23 + - + 
## 16 parent 24 1 -> 0 child 25 + - + - 
## 14 parent 23 0 -> 1 child 24 + 
## 16 parent 24 1 -> 0 child 25 + - 
## 23 parent 28 0 -> 1 child 29 + - + 
## 27 parent 30 1 -> 0 child 14 + - + - 
## 29 parent 31 0 -> 1 child 15 + - + - + 
## 1 parent 17 1 -> 0 child 18 - 
## 17 parent 25 1 -> 0 child 26 - - 
## 2 parent 18 0 -> 1 child 19 + 
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Gains et pertes de gènes

Flux de gènes avec gains et pertes

Flux de gènes

Bilan des flux de gènes avec gains et pertes