title: Variational modularity at the cell level: insights from the sperm head of the house mouse.
abstract: BACKGROUND
  Modularity is an important feature in the evolvability of organisms, since it allows the occurrence of complex adaptations at every single level of biological systems. While at the cellular level the modular organization of molecular interactions has been analyzed in detail, the phenotypic modularity (or variational modularity) of cell shape remains unexplored. The mammalian spermatozoon constitutes one of the most complex and specialized cell types found in organisms. The structural heterogeneity found in the sperm head suggests an association between its inner composition, shape and specificity of function. However, little is known about the extent of the connections between these features. Taking advantage of the house mouse sperm morphology, we analyzed the variational modularity of the sperm head by testing several hypotheses related to its structural and functional organization. Because chromosomal rearrangements can affect the genotype-phenotype map of individuals and thus modify the patterns of covariation between traits, we also evaluate the effect of Robertsonian translocations on the modularity pattern of the sperm head.
  
  
  RESULTS
  The results indicated that the house mouse sperm head is divided into three variational modules (the acrosomal, post-acrosomal and ventral spur module), which correspond to the main regions of the cytoskeletal mesh beneath the plasma membrane, i.e., the perinuclear theca. Most of the covariation is concentrated between the ventral spur and the acrosomal and post-acrosomal modules. Although the Rb fusions did not alter the main modularity pattern, they did affect the percentages of covariation between pairs of modules.
  
  
  CONCLUSIONS
  The structural heterogeneity of the cytoskeleton is responsible for the modular organization of the sperm head shape, corroborating the role that this structure has in maintaining the cell shape. The reduction in percentages of shape covariation between pairs of modules in Rb sperms suggests that chromosomal rearrangements could induce changes in the genotype-phenotype map. Nevertheless, how these variations affect sperm fertilization success is yet to be elucidated.
authors.affiliation: Departament de Biologia Animal, Biologia Vegetal i Ecologia, Facultat de Biociències, Universitat Autònoma de Barcelona, Cerdanyola del Vallès 08193, Spain. nuriaestel.medarde@uab.cat.
authors.forename: Nuria
authors.initials: N
authors.lastname: Medarde
authors.forename: Francesc
authors.initials: F
authors.lastname: Muñoz-Muñoz
authors.forename: María José
authors.initials: MJ
authors.lastname: López-Fuster
authors.forename: Jacint
authors.initials: J
authors.lastname: Ventura
mesh_terms: D000177:Acrosome / Q000648:ultrastructure; D000818:Animals; D048430:Cell Shape; D003599:Cytoskeleton / Q000737:chemistry; D005306:Fertilization; D008297:Male; D051379:Mice; D013077:Sperm Head / Q000648:ultrastructure*; D013094:Spermatozoa / Q000166:cytology*