Amphioxus molecular biology: insights into vertebrate evolution and developmental mechanisms

The cephalochordate amphioxus is the best available proxy for the last common invertebrate ancestor of the vertebrates. During the last decade, the developmental genetics of amphioxus have been extensively examined for insights into the evolutionary origin and early evolution of the vertebrates. Comparisons between expression domains of homologous genes in amphioxus and vertebrates have strengthened proposed homologies between specific body parts. Molecular genetic studies have also highlighted parallels in the developmental mechanisms of amphioxus and vertebrates. In both groups, a similar nested pattern of Hox gene expression is involved in rostrocaudal patterning of the neural tube, and homologous genes also appear to be involved in dorsoventral neural patterning. Studies of amphioxus molecular biology have also hinted that the protochordate ancestor of the vertebrates included cell populations that modified their developmental genetic pathways during early vertebrate evolution to yield definitive neural crest and neurogenic placodes. We also discuss how the application of expressed sequence tag and gene-mapping approaches to amphioxus have combined with developmental studies to advance our understanding of chordate genome evolution. We conclude by considering the potential offered by the sequencing of the amphioxus genome, which was completed in late 2004.

Complex history of a chromosomal paralogy region: insights from amphioxus aromatic amino acid hydroxylase genes and insulin-related genes

Aromatic amino acid hydroxylase (AAAH) genes and insulin-like genes form part of an extensive paralogy region shared by human chromosomes 11 and 12, thought to have arisen by tetraploidy in early vertebrate evolution. Cloning of a complementary DNA (cDNA) for an amphioxus (Branchiostoma floridae) hydroxylase gene (AmphiPAH) allowed us to investigate the ancestry of the human chromosome 11/12 paralogy region. Molecular phylogenetic evidence reveals that AmphiPAH is orthologous to vertebrate phenylalanine (PAH) genes; the implication is that all three vertebrate AAAH genes arose early in metazoan evolution, predating vertebrates. In contrast, our phylogenetic analysis of amphioxus and vertebrate insulin-related gene sequences is consistent with duplication of these genes during early chordate ancestry. The conclusion is that two tightly linked gene families on human chromosomes 11 and 12 were not duplicated coincidentally. We rationalize this paradox by invoking gene loss in the AAAH gene family and conclude that paralogous genes shared by paralogous chromosomes need not have identical evolutionary histories.

Molecular evidence from Ciona intestinalis for the evolutionary origin of vertebrate sensory placodes

Cranial sensory placodes are focused areas of the head ectoderm of vertebrates that contribute to the development of the cranial sense organs and their associated ganglia. Placodes have long been considered a key character of vertebrates, and their evolution is proposed to have been essential for the evolution of an active predatory lifestyle by early vertebrates. Despite their importance for understanding vertebrate origins, the evolutionary origin of placodes has remained obscure. Here, we use a panel of molecular markers from the Six, Eya, Pax, Dach, FoxI, COE and POUIV gene families to examine the tunicate Ciona intestinalis for evidence of structures homologous to vertebrate placodes. Our results identify two domains of Ciona ectoderm that are marked by the genetic cascade that regulates vertebrate placode formation. The first is just anterior to the brain, and we suggest this territory is equivalent to the olfactoty/adenohypophyseal placodes of vertebrates. The second is a bilateral domain adjacent to the posterior brain and includes cells fated to form the atrium and atrial siphon of adult Ciona. We show this bares most similarity to placodes fated to form the vertebrate acoustico-lateralis system. We interpret these data as support for the hypothesis that sensory placodes did not arise de novo in vertebrates, but evolved froth pre-existing specialised areas of ectoderm that contributed to sensory organs in the common ancestor of vertebrate and tunicates. Published by Elsevier Inc.

Characterisation of an amphioxus Fringe gene and the evolution of the vertebrate segmentation clock

In mouse and chick embryos, cyclic expression of lunatic fringe has an important role in the regulation of mesoderm segmentation. We have isolated a Fringe gene from the protochordate amphioxus. Amphioxus is the closest living relative of the vertebrates, and has mesoderm that is definitively segmented in a manner that is similar to, and probably homologous with, that of vertebrates. AmphiFringe is placed basal to vertebrate Fringe genes in molecular phylogenetic analyses, indicating that the duplications that formed radical-, manic- and lunatic fringe are specific to the vertebrate lineage. AmphiFringe expression was detected in the anterior neural plate of early neurulae, where it resolved into a series of segmental patches by the mid-neurulae stage. No AmphiFringe transcripts were detected in the mesoderm. Based on these observations, we propose a model depicting a successive recruitment of Fringe in the maintenance then regulation of segmentation during vertebrate evolution.

Towards a European tephrochronological framework for Termination 1 and the Early Holocene

The record of deposition of tephras in Europe and the North Atlantic during the period 18.5–8.0 ¹⁴C ka BP (the Last Termination and Early Holocene) is reviewed. Altogether, 34 tephras originating from four main volcanic provinces (Iceland, the Eifel district, the Massif Central and Italy) have been identified so far in geological sequences spanning this time–interval. Most of the records have been based, until very recently, on observations of visible layers of tephras. Here, we report on the potential for extending the areas over which some of the tephras can be traced by the search for layers of micro–tephra, which are not visible to the naked eye, and on the use of geochemical methods to correlate them with known tephra horizons. This approach has greatly extended the area in Northern Europe over which the Vedde Ash can be traced. The same potential exists in southern Europe, which is demonstrated for the first time by the discovery of a distinct layer of micro–tephra of the Neapolitan Yellow Tuff in a site in the Northern Apennines in Italy, far to the north of the occurrences of visible records of this tephra. The paper closes by considering the potential for developing a robust European tephrostratigraphy to underpin the chronology of records of the Last Termination and Early Holocene, thereby promoting a better understanding of the nature, timing and environmental effects of the abrupt climatic changes that characterized this period.

The time course of implicit affective picture processing: an eye movement study

Consistent with a negativity bias account, neuroscientific and behavioral evidence demonstrates modulation of even early sensory processes by unpleasant, potentially threat-relevant information. The aim of this research is to assess the extent to which pleasant and unpleasant visual stimuli presented extrafoveally capture attention and impact eye movement control. We report an experiment examining deviations in saccade metrics in the presence of emotional image distractors that are close to a nonemotional target. We additionally manipulate the saccade latency to test when the emotional distractor has its biggest impact on oculomotor control. The results demonstrate that saccade landing position was pulled toward unpleasant distractors, and that this pull was due to the quick saccade responses. Overall, these findings support a negativity bias account of early attentional control and call for the need to consider the time course of motivated attention when affect is implicit

The time-course of morphological constraints: Evidence from eye-movements during reading

Lexical compounds in English are constrained in that the non-head noun can be an irregular but not a regular plural (e.g. mice eater vs. *rats eater), a contrast that has been argued to derive from a morphological constraint on modifiers inside compounds. In addition, bare nouns are preferred over plural forms inside compounds (e.g. mouse eater vs. mice eater), a contrast that has been ascribed to the semantics of compounds. Measuring eyemovements during reading, this study examined how morphological and semantic information become available over time during the processing of a compound. We found that the morphological constraint affected both early and late eye-movement measures, whereas the semantic constraint for singular non-heads only affected late measures of processing. These results indicate that morphological information becomes available earlier than semantic information during the processing of compounds.

Becoming a written word: eye movements reveal order of acquisition effects following incidental exposure to new words during silent reading

We know that from mid-childhood onwards most new words are learned implicitly via reading; however, most word learning studies have taught novel items explicitly. We examined incidental word learning during reading by focusing on the well-documented finding that words which are acquired early in life are processed more quickly than those acquired later. Novel words were embedded in meaningful sentences and were presented to adult readers early (day 1) or later (day 2) during a five-day exposure phase. At test adults read the novel words in semantically neutral sentences. Participants’ eye movements were monitored throughout exposure and test. Adults also completed a surprise memory test in which they had to match each novel word with its definition. Results showed a decrease in reading times for all novel words over exposure, and significantly longer total reading times at test for early than late novel words. Early-presented novel words were also remembered better in the offline test. Our results show that order of presentation influences processing time early in the course of acquiring a new word, consistent with partial and incremental growth in knowledge occurring as a function of an individual’s experience with each word.