The evolution of modern eukaryotic phytoplankton.

The community structure and ecological function of contemporary marine ecosystems are critically dependent on eukaryotic phytoplankton. Although numerically inferior to cyanobacteria, these organisms are responsible for the majority of the flux of organic matter to higher trophic levels and the ocean interior. Photosynthetic eukaryotes evolved more than 1.5 billion years ago in the Proterozoic oceans. However, it was not until the Mesozoic Era (251 to 65 million years ago) that the three principal phytoplankton clades that would come to dominate the modern seas rose to ecological prominence. In contrast to their pioneering predecessors, the dinoflagellates, coccolithophores, and diatoms all contain plastids derived from an ancestral red alga by secondary symbiosis. Here we examine the geological, geochemical, and biological processes that contributed to the rise of these three, distantly related, phytoplankton groups.

The collapse of the two-kingdom system, the rise of protistology and the founding of the International Society for Evolutionary Protistology (ISEP).

This paper provides a brief summary of the rise and acceptance of protistology as a modern, realistic approach to the evolutionary relationships and classification of unicellular eukaryotic organisms as well as the origins of the multicellular groups. The apparent reasons for the renaissance of this 19th-century concept in the 1970s are reviewed, with electron microscopy considered to be the key factor, strongly reinforced by molecular phylogenetic studies in the 1980s and 1990s. The foundation of the International Society for Evolutionary Protistology in 1975 accompanied this major alteration in the view of biological diversity. The current status of protistology relative to protozoology and phycology is discussed.

Multiple protein phylogenies show that Oxyrrhis marina and Perkinsus marinus are early branches of the dinoflagellate lineage.

Oxyrrhis marina and Perkinsus marinus are two alveolate species of key taxonomic position with respect to the divergence of apicomplexans and dinoflagellates. New sequences from Oxyrrhis, Perkinsus and a number of dinoflagellates were added to datasets of small-subunit (SSU) rRNA, actin, alpha-tubulin and beta-tubulin sequences, as well as to a combined dataset of all three protein-coding genes, and phylogenetic trees were inferred. The parasitic Perkinsus marinus branches at the base of the dinoflagellate clade with high support in most of the individual gene trees and in the combined analysis, strongly confirming the position originally suggested in previous SSU rRNA and actin phylogenies. The SSU rRNA from Oxyrrhis marina is extremely divergent, and it typically branches with members of the Gonyaulacales, a dinoflagellate order where SSU rRNA sequences are also divergent. Conversely, none of the three protein-coding genes of Oxyrrhis is noticeably divergent and, in trees based on all three proteins individually and in combination, Oxyrrhis branches at the base of the dinoflagellate clade, typically with high bootstrap support. In some trees, Oxyrrhis and Perkinsus are sisters, but most analyses indicate that Perkinsus diverged prior to Oxyrrhis. Morphological characters have previously pointed to Oxyrrhis as an early branch in the dinoflagellate lineage; our data support this suggestion and significantly bolster the molecular data that support a relationship between Perkinsus and dinoflagellates. Together, these two organisms can be instrumental in reconstructing the early evolution of dinoflagellates and apicomplexans by helping to reveal aspects of the ancestors of both groups.

Dinoflagellate nuclear SSU rRNA phylogeny suggests multiple plastid losses and replacements.

Dinoflagellates are a trophically diverse group of protists with photosynthetic and non-photosynthetic members that appears to incorporate and lose endosymbionts relatively easily. To trace the gain and loss of plastids in dinoflagellates, we have sequenced the nuclear small subunit rRNA gene of 28 photosynthetic and four non-photosynthetic species, and produced phylogenetic trees with a total of 81 dinoflagellate sequences. Patterns of plastid gain, loss, and replacement were plotted onto this phylogeny. With the exception of the apparently early-diverging Syndiniales and Noctilucales, all non-photosynthetic dinoflagellates are very likely to have had photosynthetic ancestors with peridinin-containing plastids. The same is true for all dinoflagellates with plastids other than the peridinin-containing plastid: their ancestors have replaced one type of plastid for another, in some cases most likely through a non-photosynthetic intermediate. Eight independent instances of plastid loss and three of replacement can be inferred from existing data, but as more non-photosynthetic lineages are characterized these numbers will surely grow.

Are Pyrodinium blooms in the Southeast Asian region recurring and spreading? A view at the end of the millennium.

Pyrodinium bahamense (var. compressum) has been the only dinoflagellate species that has caused major public health and economic problems in the Southeast Asian region for more than 2 decades now. It produces saxitoxin, a suite of toxins that cause Paralytic Shellfish Poisoning (PSP). A serious toxicological problem affecting many countries of the world, mild cases of this poisoning can occur within 30 minutes while in extreme cases, death through respiratory paralysis may occur within 2-24 hrs of ingestion of intoxicated shellfish. Blooms of the organism have been reported in Malaysia, Brunei Darussalam, the Philippines and Indonesia. The ASEAN-Canada Red Tide Network has recorded 31 blooms of the organism in 26 areas since 1976 when it first occurred in Sabah, Malaysia. As of 1999, the most hard hit country has been the Philippines which has the greatest number of areas affected (18) and highest number of Paralytic Shellfish Poisoning (PSP) cases (about 1995). Malaysia has reported a total of 609 PSP cases and 44 deaths while Brunei has recorded 14 PSP cases and no fatalities. Indonesia, on the other hand has a record of 427 PSP cases and 17 deaths. Studies on ecological/environmental impacts of these blooms have not been done in the region. Estimates of economic impacts have shown that the loss could be up to USD 300,000 day-1. Most of the data and information useful for understanding Pyrodinium bloom dynamics have come from harmful/toxic algal monitoring and research that have developed to different degrees in the various countries in the region affected by the organism's bloom. Regional collaborative research and monitoring efforts can help harmonize local data sets and ensure their quality and availability for comparative analysis and modeling. Temporal patterns of the blooms at local and regional scales and possible signals and trends in the occurrence/recurrence and spread of Pyrodinium blooms could be investigated. Existing descriptive and simple predictive models of Pyrodinium blooms can be improved and refined to help in the management of the wild harvest and aquaculture of shellfish in a region where the people are dependent on these resources for their daily food sustainance and livelihood.

Chemical and biological evidence links microcystins to salmon 'netpen liver disease'.

Evidence is presented that links microcystins to a severe liver disease that occurs in Atlantic salmon that are netpen-reared in coastal British Columbia. Liquid chromatography-linked protein phosphatase bioassay analysis of extracts of liver tissue taken from Atlantic salmon afflicted with netpen liver disease showed the presence of an inhibitor of protein phosphatase that was chromatographically indistinguishable from microcystin-LR. Analysis of liver tissue from healthy control fish showed a complete absence of microcystin-LR. Intraperitoneal injection of microcystin-LR into healthy Atlantic salmon re-created the pathologic changes of netpen liver disease, including diffuse necrosis and hepatic megalocytosis.

Quality of life after heart transplantation in patients assigned to double- or triple-drug therapy.

The purpose of this study was to compare the quality of life after heart transplantation for patients treated with cyclosporine and azathioprine (double therapy) versus cyclosporine, azathioprine, and corticosteroids (triple therapy). This study was based on a randomized, prospective trial and was focused on patients from ages 17 to 57 years at 1-year after transplantation. Patients who received double therapy showed advantages on 10 of 11 measures of quality of life. Significant differences were found on measures of anxiety, sexual activity, physical well-being, and financial well-being. Patients who received double-drug therapy reported a lower frequency and less distress from the side effects of immunosuppression; a higher proportion of double-drug therapy patients had returned to full-time employment. In addition double-therapy patients were better able to control body mass. These features may explain why double-drug therapy patients reported a higher quality of life, and in the long term, could be important from a cost benefit analysis.

The code within the codons.

For the first time it is shown that each of the three codon bases has a general correlation with a different, predictable amino acid property, depending on position within the codon. In addition to the previously recognized link between the mid-base and the hydrophobic-hydrophilic spectrum, we show that, with the exception of G, the first base is generally invariant within a synthetic pathway. G--coded amino acids show a different order, being found only at the head of the synthetic pathways. The redundancy of the nature of the third base has a previously unrecognised relationship with molecular weight. The bases U and A (transversions) are associated with the most sharply defined or opposite states in both the first and second position, C somewhat less so or intermediate, anf G neutral. The apparently systematic nature of these relationships has profound implications for the origin of the genetic code. It appears to be the remains of the first language of the cell, predating the tRNA/ribosome system, persisting with remarkably little change at a deeper level of organisation than the codon language.

Non-invasive aneurysm detection using digital signal processing.

The attributes and performance expectations of a relatively low cost, non-invasive, aneurysm detection system have been studied. The developed system required that bio-acoustic data be captured at the patients' eyes, conditioned, transformed into the frequency domain, and analysed in some systematic fashion. It was admitted that the signal transducers were in need of improvement. Several low cost spectral analysers were built and tested. This system transformed the patient's bio-acoustic signal into a power spectrum. Features, representative of the patient's medical state, were extracted from the power spectrum and were input to a double-linked tree structure. Together with the medical class information, the structure can be searched to find the best statistical estimate of a patient's medical status. Experiment on generated and patient data, support the validity of the chosen diagnostic scheme.

On dinoflagellate evolution.

A broad overview of the diversity of living dinoflagellates is presented in a hypothetical evolutionary context. Ultrastructural, and some physiological information is included. Five principal organizational types: prorocentroid, dinophysoid, gonyaulacoid, peridinioid and gymnodinoid, are taken to represent lineages, and the developments within each summarized. Thecal evolution is discussed party with the aid of a model developed to determine probable plate homologies in the gonyaulacoids and peridinioids. Both primitive and highly specialized features are drawn attention to, particularly with regard to the nucleus and ocelli. The parallelism between the latter and metazoan eyes is extraordinary, considering that the dinoflagellate organelles are made of subcellular components. The roles of various types of cysts within the lifecycle of dinoflagellates are discussed. The compatibility of the hypothetical events proposed here with the fossil record is briefly considered, and some indications of the phyletic position of dinoflagellates are reviewed. The conclusions summarize the principal developments that appear to have arisen within the group. The relative primitiveness of the desmokonts is affirmed. A new combination, Plectodinium miniatum (Kofoid and Swezy) comb. nov. is proposed, as well as the recognition of a new order, the Gonyaulacales ord. nov.

Symbionticism revisited: a discussion of the evolutionary impact of intracellular symbioses.

Wallin (1927) first published the notion that the fusion of bacteria with host cells was the principal source of genetic novelty for speciation. He suggested that mitochondria are transitional elements in this process. While the significance that he attributed to symbiosis now seem excessive, he was one of the first authors to be aware of the evolutionary potential of symbiotic events and his view of mitochondria may not seem strange to many cell biologist today. The most significant evolutionary development which has been attributed to intracellular symbiosis is the origin of eukaryotic cellular organization. The current status of the 'serial endosymbiosis hypothesis' is briefly review. The case for the symbiotic origin of the chloroplast, based principally on 16 S RNA oligonucleotide cataloguing, is very strong. Mitochondrial origins are more obscure but also appear to be symbiotic due to recent 18 S cataloguing from wheat embryos. The probablility of the multiple origin of some eukaryotic organelles is also examined, the processes in question being the acquisition of distinct stocks of chloroplasts from disparate photosynthetic prokaryotes and the secondary donation of organelles from degenerate eukaryotic endosymbionts to their hosts, with specific reference to the dinoflagellates Peridinium balticum, Kryptoperidinium foliaceum and the ciliate Mesodinium rubrum. It is concluded that the evolutionary potential of intracellular symbiosis ('cytobiosis': a term introduced in this paper) is great, with the best established influence being on the origin of eukaryotic chloroplasts. Together with the potential effects of viral vectors, symbiosis serves as a supplementary speciation mechanism capable of producing directed evolutionary changes. It is likely that these processes will explain some of the apparent anomalies in evolutionary rates and direction which are not readily explicable by the conventional synthetic theory of evolution.

Problems in the development of an explicit hypothetical phylogeny of the lower eukaryotes.

A semi-explicit arrangement of the lower eukaryotes is provided to serve as a basis for phyletic discussions. No single character is used to determine the position of all the groups. The tree provides no ready separation of protozoa, algae and fungi, groups assigned to these traditional assemblages being considered to be for the most part inextricably interwoven. Photosynthetic forms, whose relationships seem to be more readily discernable, are considered to have given rise repeatedly to nonphotosynthetic forms. The assumption that there are primitive "preflagellar" eukaryotes (red algae, non-flagellated fungi) is adopted. The potential value of mitochondrial features as indicators of broad affinities is emphasised, particularly in determining the probable affinities of non-photosynthetic forms, and this criterion is contra-indicative of a ciliate ancestry for the Metazoa. In the arrangement provided the distributions of chloroplast, mitochondrial and flagellar features match one another well, suggesting their probable co-evolution.

The propulsive mechanism of the dinoflagellate transverse flagellum reconsidered.

As a result of the recent discovery that the transverse flagellum of dinoflagellates does not beat helically within the cingulum it has been necessary to formulate a new model for the manner in which this flagellum can impart a forward motion to the cell. It is theorised that the contribution to forward propulsion by the transverse flagellum can only arise directly from an asymmetry of the flagellar wave. In addition we have drawn attention to topographic features of the cell, and particularly of the left sulcal list, which may augment this thrust by redirecting cingular flow down the sulcus.

Taxonomic difficulties in red tide and paralytic shellfish poison studies: the "tamarensis complex" of Gonyaulax.

The type illustrations of the dinoflagellate Gonyaulax tamarensis contain an apparent reversal of the epithecal plates. Furthermore a culture from the type locality has been found not to be toxigenic. These two features have led a recent author to doubt the appropriateness of the allocation of toxic populations in the North Atlantic to this species or a variety of it (var. excavata Braarud). The latter has been raised to the status of a distinct species but the wrong name has been applied to it (G. excavata) as, according to the rules of priority, it should be G. phoneus (Woloszynska & Conrad) nov. comb. A history of this confused situation is provided. The criteria by which other similar species are recognised are summarised. The necessity for further study on the specific distinction of these taxa is stressed. G. conjuncta has been so inadequately described or rejected. Variability in the plate pattern of a culture of G. tamarensis var. excavata from British Columbia is illustrated and its bearing on the taxonomy of the group discussed. The presence of this toxic variety on the west coast of N. America is a new record for the Pacific Ocean.