Unlike many amphibians, the paedomorphic axolotl (Ambystoma mexicanum) rarely undergoes external morphological changes indicative of metamorphosis. However, internally, some axolotl tissues undergo cryptic metamorphic changes. A previous study examined interspecific patterns of larval brain gene expression and found that these species exhibited unique temporal expression patterns that were hypothesized to be morph specific. This thesis tested this hypothesis by examining differences in brain gene expression between juvenile (JUV), paedomorphic (PAED), and metamorphic (MET) axolotls. I identified 828 genes that were expressed differently between JUV, PAED, and MET. Expression estimates from JUV were compared to estimates from PAED and MET brains to identify genes that changed significantly during development. Genes that showed statistically equivalent expression changes across MET and PAED brains provide a glimpse at aging and maturation in an amphibian. The genes that showed statistically different expression estimates between metamorphic and paedomorphic brains provide new functional insights into the maintenance and regulation of paedomorphosis. For genes that were not commonly regulated due to aging, paedomorphs exhibited greater transcriptional similarity to juvenile than metamorphs did to juvenile. Overall, gene expression differences between metamorphic and paedomorphic development exhibit a mosaic pattern of expression as a function of aging and metamorphosis in axolotls.

A key question in biology is how changes in gene function or regulation produce new morphologies during evolution. The nectar spur is an evolutionarily labile structure known to influence speciation in a broad range of angiosperm taxa. Here, the genetic basis of nectar spur development, and the evolution of differences in nectar spur morphology, is investigated in Linaria vulgaris and two closely related species of orchid, the primitively longer-spurred Dactylorhiza fuchsii, and more derived short-spurred D. viridis (Orchidinae, Orchidaceae). Despite considerable morphological and phylogenetic differences, nectar spur ontogeny is fundamentally similar in each of the study species, proceeding from an abaxial bulge formed on the ventral petal relatively late in petal morphogenesis. However, spur development is progenetically curtailed in the short-spurred orchid D. viridis. In each case spur development involves class 1 KNOTTED1-like homeobox (KNOX) proteins. KNOX gene expression is not restricted to the spur-bearing petal, indicating that additional components are required to define nectar spur position, e.g. canonical ABC genes, determinants of floral zygomorphy, and additional (currently unknown) factors. However, constitutive expression of class 1 KNOX proteins in transgenic tobacco produces flowers with ectopic outgrowths on the petals, indicating that KNOX proteins alone are, to some degree, capable of inducing structures similar to nectar spurs in a heterologous host. Interestingly, KNOX gene expression is high in the ovary of all study taxa, suggesting that KNOX proteins may also have been involved in the evolution of this key angiosperm feature. Although principally involved in maintaining indeterminacy in the shoot apical meristem (SAM), members of the KNOX gene family have been co-opted in the evolution and development of compound leaves where they suppress differentiation and extend the morphogenetic potential of the leaf. A similar model is presented here to explain the role of KNOX proteins in nectar spur development. Co-option of KNOX gene expression to the maturing perianth delays cellular differentiation, facilitating the development of the nectar spur but requiring additional, unknown factors, to determine nectar spur fate. As facilitators of nectar spur development, changes in the spatio-temporal patterns of KNOX gene expression may alter the potential for nectar spur development and explain the critical length differences observed between the orchids D. fuchsii and D. viridis (and among other angiosperm taxa). Taken together, the available data indicate that KNOX genes confer a meristematic state upon plant tissues in a variety of morphogenetic contexts, making the gene family a potentially versatile tool to mediate a wide variety of evolutionary transformations.

Lianas de Bignoniaceae são reconhecidas por apresentarem uma variação cambial em seus caules, que promove a formação de cunhas de floema que interrompem o xilema. Uma grande diversidade de formas anatômicas foram descritas para esta variação, assim como o floema resultante dela também foi descrito como sendo distinto do floema normal presente concomitantemente nestes caules. Entretanto, nada se sabe sobre a origem, evolução e diversificação das características anatômicas neste grupo. Por essa razão, o presente estudo teve como objetivo uma análise anatômica dos caules de Bignonieae num contexto filogenético, com o intuito de lançar hipóteses para a evolução da diversidade anatômica na tribo. Para tanto, foi realizada uma análise anatômica caulinar de 54 espécies de Bignonieae, representantes dos 21 gêneros atualmente reconhecidos. Nossos resultados apontam que, não obstante a grande diversidade anatômica presente nos caules de Bignonieae, todas compartilham estágios comuns de desenvolvimento, seguidos de adições terminais que promoveram o aumento da complexidade em seus caules. Além disso, vimos que as diferenças entre o floema normal e o variante tem aumentado ao longo da evolução e está presente em todos os tipos celulares do floema. Vimos ainda que o floema secundário em Bignonieae evolui em direções opostas em diferentes linhagens da tribo, evidenciando que a evolução do floema não segue uma única direção, mas várias. Por fim, este estudo demonstra que análises anatômicas dentro de um contexto filogenético são primordiais por permitirem um maior entendimento dos processos que promoveram a evolução e diversificação dos grupos. / Lianas in Bignoniaceae are well known for presenting a cambial variant in their stems, which develops into phloem wedges that deep furrows the xylem. An enormous diversity of anatomical forms were described as resulting from this cambial variant, as well as the phloem produced by the cambial variant was described as being distinct from the regular phloem concurrently present in these stems. However, nothing is known about the origin, evolution, and diversification of the anatomical traits in this group. Therefore, the present study aimed to provide an anatomical analysis of the stems of Bignonieae (Bignoniaceae) within a phylogenetic framework, in order to address questions on the evolution of anatomical diversity in this tribe. For that reason, here we analyzed the stems of 54 species of Bignonieae, representative of the 21 genera currently known for the tribe. Our results show that, despite the great anatomical diversity present in the stems of Bignonieae, all of them share common developmental stages, which are then followed by subsequent terminal additions that are though to have promoted an augment in the complexity of these stems. Furthermore, our results indicate that the differences found between regular and variant phloem is increasing along time and is present in all cell types of the phloem. Moreover, we found that the secondary phloem in Bignonieae is evolving in opposite directions in distinct lineages of the tribe, evidencing that the evolution of the phloem is not constraint to a single line of specialization. In conclusion, this study demonstrates the importance of anatomical analyses within a phylogenetic framework, allowing for the detection of the processes that have been involved in the evolution and diversification of plant groups.

Lianas de Bignoniaceae são reconhecidas por apresentarem uma variação cambial em seus caules, que promove a formação de cunhas de floema que interrompem o xilema. Uma grande diversidade de formas anatômicas foram descritas para esta variação, assim como o floema resultante dela também foi descrito como sendo distinto do floema normal presente concomitantemente nestes caules. Entretanto, nada se sabe sobre a origem, evolução e diversificação das características anatômicas neste grupo. Por essa razão, o presente estudo teve como objetivo uma análise anatômica dos caules de Bignonieae num contexto filogenético, com o intuito de lançar hipóteses para a evolução da diversidade anatômica na tribo. Para tanto, foi realizada uma análise anatômica caulinar de 54 espécies de Bignonieae, representantes dos 21 gêneros atualmente reconhecidos. Nossos resultados apontam que, não obstante a grande diversidade anatômica presente nos caules de Bignonieae, todas compartilham estágios comuns de desenvolvimento, seguidos de adições terminais que promoveram o aumento da complexidade em seus caules. Além disso, vimos que as diferenças entre o floema normal e o variante tem aumentado ao longo da evolução e está presente em todos os tipos celulares do floema. Vimos ainda que o floema secundário em Bignonieae evolui em direções opostas em diferentes linhagens da tribo, evidenciando que a evolução do floema não segue uma única direção, mas várias. Por fim, este estudo demonstra que análises anatômicas dentro de um contexto filogenético são primordiais por permitirem um maior entendimento dos processos que promoveram a evolução e diversificação dos grupos. / Lianas in Bignoniaceae are well known for presenting a cambial variant in their stems, which develops into phloem wedges that deep furrows the xylem. An enormous diversity of anatomical forms were described as resulting from this cambial variant, as well as the phloem produced by the cambial variant was described as being distinct from the regular phloem concurrently present in these stems. However, nothing is known about the origin, evolution, and diversification of the anatomical traits in this group. Therefore, the present study aimed to provide an anatomical analysis of the stems of Bignonieae (Bignoniaceae) within a phylogenetic framework, in order to address questions on the evolution of anatomical diversity in this tribe. For that reason, here we analyzed the stems of 54 species of Bignonieae, representative of the 21 genera currently known for the tribe. Our results show that, despite the great anatomical diversity present in the stems of Bignonieae, all of them share common developmental stages, which are then followed by subsequent terminal additions that are though to have promoted an augment in the complexity of these stems. Furthermore, our results indicate that the differences found between regular and variant phloem is increasing along time and is present in all cell types of the phloem. Moreover, we found that the secondary phloem in Bignonieae is evolving in opposite directions in distinct lineages of the tribe, evidencing that the evolution of the phloem is not constraint to a single line of specialization. In conclusion, this study demonstrates the importance of anatomical analyses within a phylogenetic framework, allowing for the detection of the processes that have been involved in the evolution and diversification of plant groups.