Biomedical subjects
Publications and source records attributed to Alessandro Minelli.
Simple treatments with hormones could unlock the expression of complex phenotypes not known to occur in nature. Using this method, Kim et al. recently obtained males from all-female populations of water fleas. The novel characters revealed by this work can be used in taxonomic identification and phylogenetic inference. Additionally, these "resurrected" males offer insights into the conservation of traits that are not exposed to natural selection.
The traditional framework for the description of arthropod development takes the molt-to-molt interval as the fundamental unit of periodization, which is similar to the morphological picture of the main body axis as a series of segments. Developmental time is described as the subdivision into a few major stages of one or more instars each, which is similar to the subdivision of the main body axis into regions of one to many segments each. Parallel to recent criticisms to the segment as the fundamental building block of arthropod anatomy, we argue that, while a firm subdivision of development in stages is useful for describing arthropod ontogeny, this is limiting as a starting point for studying its evolution. Evolutionary change affects the association between different developmental processes, some of which are continuous in time whereas others are linked to the molting cycle. Events occurring but once in life (hatching; first achieving sexual maturity) are traditionally used to establish boundaries between major units of arthropod developmental time, but these boundaries are quite labile. The presence of embryonic molts, the 'gray zone' of development accompanying hatching (with the frequent delivery of an immature whose qualification as 'free-embryo' or ordinary postembryonic stage is arbitrary), and the frequent decoupling of growth and molting suggest a different view. Beyond the simple comparison of developmental schedules in terms of heterochrony, the flexible canvas we suggest for the analysis of arthropod development opens new vistas into its evolution. Examples are provided as to the origin of holometaboly and hypermetaboly within the insects.
The origin of evolutionary novelties is a central topic in evolutionary developmental biology (evo-devo) studies. In any new feature, there is a conserved component that is either structural or related to the underlying genetic control, but it is not always obvious what is really new and what is conserved. Nevertheless, disentangling this blending of old and new features is basic to understanding mechanisms of evolutionary change. The origin of arthropod appendages illustrates the complexity in tracing the origin of evolutionary novelties. At the base of the lineage, the main body axis was already segmented and antero-posteriorly patterned, and the genetic tool kit required to form lateral outgrowths was already available. The novelty was possibly the developmental decision to "read" the available axial information and to exploit it for differentiating segmentally patterned and axially segmented appendages. Some important novelties bridge the gap between what have been traditionally distinguished as systemic and local changes. For example, the origin of the body cavities evolved by several animal groups may have been initiated by simple changes in cell-to-cell adhesive properties. Any possible change in an existing ontogenetic pathway has the potential to generate novelties.
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Recent advances in molecular phylogenetics are continuously changing our perception of the phylogenetic relationships among the main arthropod lineages: crustaceans, hexapods, chelicerates, and myriapods. Besides the intrinsic interest in unraveling the evolution of the largest animal phylum, these studies are basic to an understanding of one of the major transitions in animal evolution-i.e., the conquest of land with all its associated structural and functional adaptations. Myriapods have been traditionally considered the closest relatives of hexapods, thus implying only one origin of terrestriality for the tracheate lineage, but this view is now challenged by molecular evidence. Sequence data available to date for centipedes and millipedes are very limited, and the taxon sampling is strongly biased. The most critical gap was the scutigeromorph centipedes, which are the sister group to all remaining Chilopoda from which they probably diverged in the Silurian if not earlier. We obtained the first complete mitochondrial sequence for a representative of this clade, the house centipede. In our phylogenetic analyses of the protein-coding genes in this mitochondrial genome, along with 16 further ones representing the other major arthropod clades plus two outgroups, the myriapods formed a clade with the chelicerates. This implies that water-to-land transition occurred at least three times (hexapods, myriapods, arachnids) during the evolution of the Arthropoda. In addition, in contrast to all previous studies, our best supported topologies favor paraphyly of the myriapods with respect to the chelicerates. This would increase to four the main events of land colonization in arthropods (once for centipedes, once for millipedes).
We describe the complete mitochondrial genome of the house centipede Scutigera coleoptrata. Its gene order is unique among characterized arthropod mitochondrial genomes. Comparison to the gene order in the horseshoe crab mtDNA implies 10 or more translocations. By extending comparisons to 30 arthropod mitochondrial genomes plus two outgroups, we identify two different patterns of gene order change. The first, only affecting position and orientation of tRNAs, is much more frequent than the second, which also involves protein encoding and ribosomal genes. The analysis of the same data set using available algorithms for phylogenetic reconstruction based on gene order results in unreliable trees. This indicates that the current methods for analyzing gene order rearrangement are not suitable for wide-ranging phylogenetic studies.
Bilaterian animals show a diverse array of segmental patterns and segmentation processes. Differences in pattern and process emerge both in comparisons of taxa and among sets of serial structures within one animal. Diversity in developmental mechanisms of segmentation and their genetic control is reflected in the modes in which segmentation evolves, which are difficult to accommodate within the traditional concept of segments as modular building blocks. Thus, in spite of the apparent simplicity of segmental patterns, studying the evolution of segmentation requires an approach that, in an adequate comparative framework, combines the efforts of researchers of genes, cells, embryos and post-embryonic stages.
Trilobites offer the opportunity to explore postembryonic development within the fossil record of arthropod evolution. In contrast to most trilobites, the Silurian proetid Aulacopleura konincki from the Czech Republic exhibits marked variation in the mature number of thoracic segments, with five morphs with 18-22 thoracic segments. The combination of abundant articulated specimens available from a narrow stratigraphic interval and segmental intraspecific variation makes this trilobite singularly useful for studying postembryonic growth and segmentation. Trunk segmentation followed a hemianamorphic pattern, as seen in other arthropods and as characteristic of the Trilobita; during a first anamorphic phase, segments were accreted, while in the subsequent epimorphic phase, segmentation did not proceed further despite continued growth. Size increment during the anamorphic phase was targeted and followed Dyar's rule, a geometric progression typical of many arthropods. We consider alternative hypotheses for the control of the switch from anamorphic to epimorphic phases of development. Our analysis favors a scenario in which the mature number of thoracic segments was determined quite early in development rather than at a late stage in association with a critical size threshold. This study demonstrates that hypotheses concerning developmental pattern and control can be tested in organisms belonging to an extinct clade.
Many different and even contrasting notions of homology have been proposed over two centuries of comparative biology, beginning beginning with the initial reference to idealised archetypes, down to the current concepts based on derivation from a common ancestor, or on shared developmental pathways or genetic (more extensively, informational) background. Select anatomical features such as the patterns of innervation, or the expression patterns of genes putatively involved in key developmental events, e.g. the Hox genes, have been repeatedly suggested as the most reliable cues to homology. This confidence, however, rests on shaky ground and results are never certain. Recent work in comparative morphology and evolutionary developmental biology increasingly suggests the need to abandon the traditional all-or-nothing notion of homology, in favour of a more flexible, factorial or combinatorial approach. In this way it will be possible to accommodate within one broad comparative view, respectful of phylogeny and developmental biology alike, many disparate notions such as positional and special homology, serial homology and temporal serial homology. All statements of homology, however, will thus require inadequate qualification of the context specifically taken in consideration and the criteria used to address the comparison. It remains to be seen, in the near future, how far the old concept of homology, now under the burden of so many and so different notions, will still be of use to comparative biology.
Current awareness of gene expression patterns and developmental mechanisms involved in the outgrowth and patterning of animal appendages contributes to our understanding of the origin and evolution of these body parts. Nevertheless, this vision needs to be complemented by a new adequate comparative framework, in the context of a factorial notion of homology. It may even be profitable to categorize as appendages also gut diverticula, body ingrowths and 'virtual appendages' such as the eye spots on butterfly wings. Another unwarranted framework is the Cartesian co-ordinate system onto which the appendages are currently described and where it is supposed that one patterning system exists for each separate Cartesian axis. It may be justified, instead, to look for correspondences between the appendages and the main body axis of the same animal, as the latter might be the source of the growth and patterning mechanisms which gave rise to the former. This hypothesis of axis paramorphisms is contrasted with the current hypothesis of gene co-option. Recapitulationism is a common fault in current Evo-Devo perspectives concerning the origin of the appendages, in that the evolutionary origin of appendages is often expected to be the same as one of the key mechanisms involved in the ontogenetic inception of appendage formation. This unwarranted perspective is also evident in the current debate on the nature of the default arthropod appendage. Most likely, a default arthropod appendage never did exist, as the first appendages probably developed along the trunk of an animal already patterned extensively along the antero-posterior body axis.
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Here we report on a partial screen for Hox gene sequences from the geophilomorph centipede Pachymerium ferrugineum, resulting in 11 different sequences. All of these sequences could be homologized to specific Drosophila genes, yielding two representatives for the Dfd class and one each for the remaining classes. Phylogenetic analysis of these data with a broad sample of arthropod/onychophoran homologous sequences confirmed these results and provided further support for the monophyly of the Hox3/zen class. Conversely, the phylogenetic status of ftz-type genes remains uncertain. Our results complement the previous partial findings for two other centipedes (the scolopendromorph Ethmostigmus rubripes and the lithobiomorph Lithobius forficatus) and confirm the expectation that in myriapods, too, all Hox genes classes are present. This suggests that even the Chilopoda, with uniform trunk segments, have the same number of Hox genes as the more tagmatized Insecta.
Similarities in genetic control between the main body axis and its appendages have been generally explained in terms of genetic co-option. In particular, arthropod and vertebrate appendages have been explained to invoke a common ancestor already provided with patterned body outgrowths or independent recruitment in limb patterning of genes or genetic cassettes originally used for purposes other than axis patterning. An alternative explanation is that body appendages, including genitalia, are evolutionarily divergent duplicates (paramorphs) of the main body axis. However, are all metazoan limbs and genitalia homologous? The concept of body appendages as paramorphs of the main body axis eliminates the requirement for the last common ancestor of limb-bearing animals to have been provided with limbs. Moreover, the possibility for an animal to express complex organs ectopically demonstrates that positional and special homology may be ontogenetically and evolutionarily uncoupled. To assess the homology of animal genitalia, we need to take into account three different sets of mechanisms, all contributing to their positional and/or special homology and respectively involved (1) in the patterning of themain body axis, (2) in axis duplication, followed by limb patterning mechanisms diverging away from those still patterning the main body axis (axis paramorphism), and (3) in controlling the specification of sexual/genital features, which often, but not necessarily, come into play by modifying already developed and patterned body appendages. This analysis demonstrates that a combinatorial approach to homology helps disentangling phylogenetic and ontogenetic layers of homology.
When approaching the study of terminal growth and segmentation, comparative morphology provides an important guide to formulate questions. There are often problems in unambiguously identifying the axis along which we wish to study terminal (often, actually, subterminal) growth, especially when the trunk axis is posteriorly prolonged in an appendage (as with the tail of vertebrates), or when the polarity of the "external animal" is other than the polarity of the "internal animal," as in polypoid bilaterians. We cannot ignore that the rear end of the main body axis is possibly defined very early in development in some groups, for example, arthropods, whereas in others, vertebrates, for example, it is defined much later. We cannot think of segmentation as always corresponding to the sequential posterior addition of new units, thus ignoring the widespread occurrence of double segmentation. A more subtle problem is represented by the overlapping of different processes, all of them contributing to elongating the body, such as segmentation, cell proliferation, cell rearrangement, and cell growth. Within a segmented trunk, cell proliferation and differentiation may go on in parallel from as many growth points as there are groups of regularly spaced cells. The main consequence, however, is not so much to expedite elongation as to reduce the disparity of metabolic conditions, gene expression patterns and "relative age" of different body districts, otherwise possibly troublesome within the limited space of the embryo.
We report the coding sequence and embryonic expression of the four trunk Hox genes Antennapedia (Antp), Ultrabithorax (Ubx), abdominal-A (abd-A), and Abdominal-B (Abd-B) in the geophilomorph centipede Strigamia maritima. In geophilomorph centipedes, all leg-bearing segments (LBS) are generated during embryogenesis, allowing us to define expression in relation to the full extent of the forming trunk. Persistent Antp expression characterizes the maxillipedal (poison claw) segment, whereas all LBS express the three Hox genes Antp, Ubx, and abd-A. Abd-B is never detectably expressed in segmented tissue, but is restricted to a zone around the proctodaeum that contributes to the hindgut. Expression of all these Hox genes initiates in the unsegmented tissue of the blastodisc, with expression of Antp respecting a sharply defined anterior border before the appearance of morphological segmentation in the trunk. The accumulation of Hox gene transcripts is strongly modulated by the maturing segment pattern, suggesting regulatory interactions with multiple levels of the segment patterning machinery. For one of these genes, Ubx, we detect both sense and anti-sense transcripts. The anti-sense transcripts originate 3' to the Ubx coding sequence and overlap the homeobox exon; they are expressed earlier than the Ubx coding transcripts and persistently, in an axially restricted pattern comparable to but distinct from those of the Hox coding transcripts. The pattern of accumulation of Ubx sense and anti-sense transcripts is strikingly complementary, suggesting the possibility of anti-sense regulation of Ubx expression.