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Paleobiology, community ecology, and scales of ecological pattern

The fossil record provides a wealth of data on the role of regional processes and historical events in shaping biological communities over a variety of time scales. The Quaternary record with its evidence of repeated climatic change shows that both terrestrial and marine species shifted independently rather than as cohesive assemblages over scales of thousands of years. Larger scale patterns also show a strong individualistic component to taxon dynamics; assemblage stability, when it occurs, is difficult to separate from shared responses to low rates of environmental change. Nevertheless, the fossil record does suggest that some biotic interactions influence large-scale ecological and evolutionary patterns, albeit in more diffuse and protracted fashions than those generally studied by community ecologists. These include: (1) the resistance by incumbents to the establishment of new or invading taxa, with episodes of explosive diversification often appearing contingent on the removal of incumbents at extinction events; (2) steady states of within-habitat and global diversity at longer time scales (10(7)-l0(8) yr), despite enormous turnover of taxa; and (3) morphological and biogeographic responses to increased intensities of predation and substratum disturbance over similarly long time scales. The behavior of species and communities over the array of temporal and spatial scales in the fossil record takes on additional significance for framing conservation strategies, and for understanding recovery of species, lineages, and communities from environmental changes.

NASA Discipline Exobiology↗

Thinking of biology: asteroid impacts, microbes, and the cooling of the atmosphere

The authors examine the cooling of the Earth's surface from 3.75 to 1 billion years ago. Three effects of the bombardment of Earth by asteroids and comets that may have delayed surface cooling include time to form continents, volatilization of carbonate rocks which released carbon dioxide into the atmosphere, and inability of microbes to inhabit land masses during large impact events. Continental microbes may have helped reduce high temperatures from 3.75 to 3.5 billion years ago. If so, the evolutionary sequence of microbes is proposed to be anaerobic heterotrophs, chemoautotrophs, and then photoautotrophs.

Non-NASA Center↗

Adaptation, plant evolution, and the fossil record

The importance of adaptation in determining patterns of evolution has become an important focus of debate in evolutionary biology. As it pertains to paleobotany, the issue is whether or not adaptive evolution mediated by natural selection is sufficient to explain the stratigraphic distributions of taxa and character states observed in the plant fossil record. One means of addressing this question is the functional evaluation of stratigraphic series of plant organs set in the context of paleoenvironmental change and temporal patterns of floral composition within environments. For certain organ systems, quantitative estimates of biophysical performance can be made on the basis of structures preserved in the fossil record. Performance estimates for plants separated in time or space can be compared directly. Implicit in different hypotheses of the forces that shape the evolutionary record (e.g. adaptation, mass extinction, rapid environmental change, chance) are predictions about stratigraphic and paleoenvironmental trends in the efficacy of functional performance. Existing data suggest that following the evolution of a significant structural innovation, adaptation for improved functional performance can be a major determinant of evolutionary changes in plants; however, there are structural and development limits to functional improvement, and once these are reached, the structure in question may no longer figure strongly in selection until and unless a new innovation evolves. The Silurian-Devonian paleobotanical record is consistent with the hypothesis that the succession of lowland floodplain dominants preserved in the fossil record of this interval was determined principally by the repeated evolution of new taxa that rose to ecological importance because of competitive advantages conferred by improved biophysical performance. This does not seem to be equally true for Carboniferous-Jurassic dominants of swamp and lowland floodplain environments. In these cases, environmental disruption appears to have been a major factor in shaping the fossil record. This does not mean that continuing adaptation was not important during this interval, but it may indicate that adaptive evolution was strongest in environments other than those best represented in the paleobotanical record.

NASA Discipline Exobiology↗

Antitropical distribution and evolution in the Indo-West Pacific Ocean

Antitropical distributions of continental shelf, Indo-West Pacific species are probably not due to transgression of the tropics during the glacial periods, isothermic submergence, island integration, rising Neogene temperatures, or the Mesozoic dispersal of fragments from a Pacific continental mass. Characteristics of common antitropical patterns, plus information from systematic works on a variety on a variety of animal and plant groups, indicate that the long discarded "relict theory" of Theel (1885) appears to best fit the evidence, for it provides a mechanism whereby antitropical distribution may be brought about. The relict theory is compatible with the concept that the East Indies part of the Indo-West Pacific has been functioning as a center of evolutionary origin. It suggests that antitropical and associated disjunct patterns are produced as an older species, that has spread out to occupy a broad range, loses ground and gradually becomes supplanted by a younger species that had subsequently evolved in the East Indies. As this process goes on, the older species becomes restricted to a few isolated localities on the fringe of its original range. These isolates are often found to the north and south of the equatorial region but may include relict populations at the western edge of the Indian Ocean.

NASA Discipline Exobiology↗

Extinction from a paleontological perspective

Extinction of widespread species is common in evolutionary time (millions of years) but rare in ecological time (hundreds or thousands of years). In the fossil record, there appears to be a smooth continuum between background and mass extinction; and the clustering of extinctions at mass extinctions cannot be explained by the chance coincidence of independent events. Although some extinction is selective, much is apparently random in that survivors have no recognizable superiority over victims. Extinction certainly plays an important role in evolution, but whether it is constructive or destructive has not yet been determined.

Review↗

The origin and evolution of oxygenic photosynthesis

The evolutionary developments that led to the ability of photosynthetic organisms to oxidize water to molecular oxygen are discussed. Two major changes from a more primitive non-oxygen-evolving reaction center are required: a charge-accumulating system and a reaction center pigment with a greater oxidizing potential. Intermediate stages are proposed in which hydrogen peroxide was oxidized by the reaction center, and an intermediate pigment, similar to chlorophyll d, was present.

Review, Tutorial↗

The universal ancestor

A genetic annealing model for the universal ancestor of all extant life is presented; the name of the model derives from its resemblance to physical annealing. The scenario pictured starts when "genetic temperatures" were very high, cellular entities (progenotes) were very simple, and information processing systems were inaccurate. Initially, both mutation rate and lateral gene transfer levels were elevated. The latter was pandemic and pervasive to the extent that it, not vertical inheritance, defined the evolutionary dynamic. As increasingly complex and precise biological structures and processes evolved, both the mutation rate and the scope and level of lateral gene transfer, i.e., evolutionary temperature, dropped, and the evolutionary dynamic gradually became that characteristic of modern cells. The various subsystems of the cell "crystallized," i.e., became refractory to lateral gene transfer, at different stages of "cooling," with the translation apparatus probably crystallizing first. Organismal lineages, and so organisms as we know them, did not exist at these early stages. The universal phylogenetic tree, therefore, is not an organismal tree at its base but gradually becomes one as its peripheral branchings emerge. The universal ancestor is not a discrete entity. It is, rather, a diverse community of cells that survives and evolves as a biological unit. This communal ancestor has a physical history but not a genealogical one. Over time, this ancestor refined into a smaller number of increasingly complex cell types with the ancestors of the three primary groupings of organisms arising as a result.

NASA Discipline Exobiology↗

Microfossils from the Neoarchean Campbell Group, Griqualand West Sequence of the Transvaal Supergroup, and their paleoenvironmental and evolutionary implications

The oldest filament- and colonial coccoid-containing microbial fossil assemblage now known is described here from drill core samples of stromatolitic cherty limestones of the Neoarchean, approximately 2600-Ma-old Campbell Group (Ghaap Plateau Dolomite, Lime Acres Member) obtained at Lime Acres, northern Cape Province, South Africa. The assemblage is biologically diverse, including entophysalidacean (Eoentophysalis sp.), probable chroococcacean (unnamed colonial coccoids), and oscillatoriacean cyanobacteria (Eomycetopsis cf. filiformis, and Siphonophycus transvaalensis), as well as filamentous fossil bacteria (Archaeotrichion sp.); filamentous possible microfossils (unnamed hematitic filaments) also occur. The Campbell Group microorganisms contributed to the formation of stratiform and domical to columnar stromatolitic reefs in shallow subtidal to intertidal environments of the Transvaal intracratonic sea. Although only moderately to poorly preserved, they provide new evidence regarding the paleoenvironmental setting of the Campbell Group sediments, extend the known time-range of entophysalidacean cyanobacteria by more than 400 million years, substantiate the antiquity and role in stromatolite formation of Archean oscillatoriacean cyanobacteria, and document the exceedingly slow (hypobradytelic) evolutionary rate characteristic of this early evolving prokaryotic lineage.

Non-NASA Center↗

Orthologs, paralogs and genome comparisons

During the past decade, ancient gene duplications were recognized as one of the main forces in the generation of diverse gene families and the creation of new functional capabilities. New tools developed to search data banks for homologous sequences, and an increased availability of reliable three-dimensional structural information led to the recognition that proteins with diverse functions can belong to the same superfamily. Analyses of the evolution of these superfamilies promises to provide insights into early evolution but are complicated by several important evolutionary processes. Horizontal transfer of genes can lead to a vertical spread of innovations among organisms, therefore finding a certain property in some descendants of an ancestor does not guarantee that it was present in that ancestor. Complete or partial gene conversion between duplicated genes can yield phylogenetic trees with several, apparently independent gene duplications, suggesting an often surprising parallelism in the evolution of independent lineages. Additionally, the breakup of domains within a protein and the fusion of domains into multifunctional proteins makes the delineation of superfamilies a task that remains difficult to automate.

Non-NASA Center↗

Developmental evolutionary biology of the vertebrate ear: conserving mechanoelectric transduction and developmental pathways in diverging morphologies

This brief overview shows that a start has been made to molecularly dissect vertebrate ear development and its evolutionary conservation to the development of the insect hearing organ. However, neither the patterning process of the ear nor the patterning process of insect sensory organs is sufficiently known at the moment to provide more than a first glimpse. Moreover, hardly anything is known about otocyst development of the cephalopod molluscs, another triploblast lineage that evolved complex 'ears'. We hope that the apparent conserved functional and cellular components present in the ciliated sensory neurons/hair cells will also be found in the genes required for vertebrate ear and insect sensory organ morphogenesis (Fig. 3). Likewise, we expect that homologous pre-patterning genes will soon be identified for the non-sensory cell development, which is more than a blocking of neuronal development through the Delta/Notch signaling system. Generation of the apparently unique ear could thus represent a multiplication of non-sensory cells by asymmetric and symmetric divisions as well as modification of existing patterning process by implementing novel developmental modules. In the final analysis, the vertebrate ear may come about by increasing the level of gene interactions in an already existing and highly conserved interactive cascade of bHLH genes. Since this was apparently achieved in all three lineages of triploblasts independently (Fig. 3), we now need to understand how much of the morphogenetic cascades are equally conserved across phyla to generate complex ears. The existing mutations in humans and mice may be able to point the direction of future research to understand the development of specific cell types and morphologies in the formation of complex arthropod, cephalopod, and vertebrate 'ears'.

Non-NASA Center↗

Monolignol radical-radical coupling networks in western red cedar and Arabidopsis and their evolutionary implications

The discovery of a nine-member multigene dirigent family involved in control of monolignol radical-radical coupling in the ancient gymnosperm, western red cedar, suggested that a complex multidimensional network had evolved to regulate such processes in vascular plants. Accordingly, in this study, the corresponding promoter regions for each dirigent multigene member were obtained by genome-walking, with Arabidopsis being subsequently transformed to express each promoter fused to the beta-glucuronidase (GUS) reporter gene. It was found that each component gene of the proposed network is apparently differentially expressed in individual tissues, organs and cells at all stages of plant growth and development. The data so obtained thus further support the hypothesis that a sophisticated monolignol radical-radical coupling network exists in plants which has been highly conserved throughout vascular plant evolution.

NASA Discipline Plant Biology↗

A Vendian-Cambrian boundary succession from the northwestern margin of the Siberian Platform: stratigraphy, palaeontology, chemostratigraphy and correlation

Siberia contains several key reference sections for studies of biological and environmental evolution across the Proterozoic-Phanerozoic transition. The Platonovskaya Formation, exposed in the Turukhansk region of western Siberia, is an uppermost Proterozoic to Cambrian succession whose trace and body fossils place broad limits on the age of deposition, but do not permit detailed correlation with boundary successions elsewhere. In contrast, a striking negative carbon isotopic excursion in the lower part of the Platonovskaya Formation permits precise chemostratigraphic correlation with upper-most Yudomian successions in Siberia, and possibly worldwide. In addition to providing a tool for correlation, the isotopic excursion preserved in the Platonovskaya and contemporaneous successions documents a major biogeochemical event, likely involving the world ocean. The excursion coincides with the palaeontological breakpoint between Ediacaran- and Cambrian-style assemblages, suggesting a role for biogeochemical change in evolutionary events near the Proterozoic Cambrian boundary.

Non-NASA Center↗

Disparate rates, differing fates: tempo and mode of evolution changed from the Precambrian to the Phanerozoic

Over the past quarter century, detailed genus- and species-level similarities in cellular morphology between described taxa of Precambrian microfossils and extant cyanobacteria have been noted and regarded as biologically and taxonomically significant by numerous workers world-wide. Such similarities are particularly well documented for members of the Oscillatoriaceae and Chroococcaceae, the two most abundant and widespread Precambrian cyanobacterial families. For species of two additional families, the Entophysalidaceae and Pleurocapsaceae, species-level morphologic similarities are supported by in-depth fossil-modern comparisons of environment, taphonomy, development, and behavior. Morphologically and probably physiologically as well, such cyanobacterial "living fossils" have exhibited an extraordinarily slow (hypobradytelic) rate of evolutionary change, evidently a result of the broad ecologic tolerance characteristic of many members of the group and a striking example of G. G. Simpson's [Simpson, G.G. (1944) Tempo and Mode in Evolution (Columbia Univ. Press, New York)] "rule of the survival of the relatively unspecialized." In both tempo and mode of evolution, much of the Precambrian history of life--that dominated by microscopic cyanobacteria and related prokaryotes--appears to have differed markedly from the more recent Phanerozoic evolution megascopic, horotelic, adaptationally specialized eukaryotes.

Review↗

Evolutionary dynamics of plants and animals: a comparative approach

Patterns of longevity and rate of appearance of taxa in the fossil record indicate a different evolutionary dynamic between land plants and marine invertebrates. Among marine invertebrates, rates of taxonomic turnover declined through the Phanerozoic, with increasingly extinction-resistant, long-lived, clades coming to dominate. Among terrestrial vascular plants, rates of turnover increased through the Phanerozoic, with short-lived, extinction-prone clades coming to dominate from the Devonian to the present. Terrestrial vertebrates appear to approximate the marine invertebrate pattern more closely than the plant record. We identify two features which individually or jointly may have influenced this distinction. First, land plants continuously invaded stressful environments during their evolution, while marine invertebrates and terrestrial vertebrates did not. Second, the relative structural simplicity and indeterminate mode of plant growth vs. the relative structural complexity and determinate mode of animal growth may have influenced the timing of major clade origin in the two groups.

Review↗

Carbonate deposition during the late Proterozoic Era: an example from Spitsbergen

Carbonate sediments reflect the physico-chemical and biological circumstances of their formation; thus, features of limestones and dolomites may provide insights into both environmental and evolutionary change through geological time. The Upper Proterozoic (approx 800-700 Ma) Akademikerbreen Group, Spitsbergen, comprises 2000 m of carbonates, with only minor intercalations of quartz arenite and shale. Although Proterozoic carbonates are often seen as predominantly dolomitic, the Akademikerbreen Group is about 45 percent limestone. Stromatolites are conspicuous in outcrop but constitute only 25 percent of the total section. Micrites and coarser intraclastic carbonates derived mainly from micritric precursors comprise 60 percent of the group, while oolites make up the remaining 15 percent. Distinctive sedimentary features of the group include giant (up to 16 mm) ooids, very early diagenetic calcite nodules and cements, micrites containing subaqueous shrinkage cracks filled with equant microspar cement, and strong 13C enrichment in both carbonates and co-occurring organic matter. The principal features of Akademikerbreen carbonates are widely distributed in coeval successions. However, these rocks appear to differ from older limestones and dolomites in their relative abundance of grainstones and, perhaps, micrites, as well as their paucity of tufa-like laminates and columnar or coniform stromatolites that preserve petrographic evidence of in situ precipitation as a dominant means of carbonate accretion. Upper Proterozoic carbonates also differ from Paleozoic accumulations, but the transition is not abrupt. Most changes accompanying the Proterozoic/Phanerozoic transition can be interpreted in terms of the consequences rather than the causes of metazoan and metaphyte evolution, including the evolution of biomineralization. Carbonate sedimentology reinforces data from other sources which indicate the last 200 to 300 Ma of the Proterozoic Eon was a distinctive interval of Earth history.

NASA Discipline Number 52-30↗

Oxygen and evolutionary patterns in the sea: onshore/offshore trends and recent recruitment of deep-sea faunas

Over the last 15 years a striking pattern of diversification has been documented in the fossil record of benthic marine invertebrates. Higher taxa (orders) tend to originate onshore, diversify offshore, and retreat into deep-water environments. Previous studies attribute this macroevolutionary pattern to a variety of causes, foremost among them the role of nearshore disturbance in providing opportunities for the evolution of novel forms accorded ordinal rank. Our analysis of the post-Paleozoic record of ordinal first appearances indicates that the onshore preference of ordinal origination occurred only in the Mesozoic prior to the Turonian stage of the Cretaceous, a period characterized by relatively frequent anoxic/dysoxic bottom conditions in deeper marine environments. Later, in the Cretaceous and Cenozoic, ordinal origination of benthic organisms did not occur exclusively, or even preferentially, in onshore environments. This change in environmental pattern of ordinal origination roughly correlates with Late Cretaceous: (i) decline in anoxia/dysoxia in offshore benthic environments; (ii) extinction of faunas associated with dysoxic conditions; (iii) increase in bioturbation with the expansion of deep burrowing forms into offshore environments; and (iv) offshore expansion of bryozoan diversity. We also advance a separate argument that the Cenomanian/Turonian and latest Paleocene global events eliminated much of the deep-water benthos. This requires a more recent origin of modern vent and deep-sea faunas, from shallower water refugia, than the Paleozoic or early Mesozoic origin of these faunas suggested by other workers.

Non-NASA Center↗

A model of onshore-offshore change in faunal diversity

Onshore-offshore patterns of faunal change occurred at many taxonomic scales during the Paleozoic Era, ranging from replacement of the Cambrian evolutionary fauna by the Paleozoic fauna to the environmental expansion of many orders and classes. A simple mathematical model is constructed to investigate such change. The environmental gradient across the marine shelf-slope is treated as a linear array of discrete habitats, each of which holds a set number of species, as observed in the fossil record. During any interval of time, some portion of the species in each habitat becomes extinct by background processes, with rates of extinction varying among both clades and habitats, as also observed in the record. After extinction, species are replaced from within the habitat and from immediately adjacent habitats, with proportions dependent on surviving species. This model leads to the prediction that extinction-resistant clades will always diversify at the expense of extinction-prone clades. But if extinction intensity is highest in nearshore habitats, extinction-resistant clades will expand preferentially in the onshore direction, build up diversity there, and then diversify outward toward the offshore. Thus, onshore-offshore patterns of diversification may be the expectation for faunal change quite independently of whether or not clades originate onshore. When the model is parameterized for Paleozoic trilobites and brachiopods, numerical solutions exhibit both a pattern of faunal change and a time span for diversification similar to that seen in the fossil record. They also generate structure similar to that seen in global diversification, including logistic patterns of growth, declining origination but constant extinction within clades through time, and declining overall extinction across clades through time.

NASA Discipline Number 52-40↗

On the early emergence of reverse transcription: theoretical basis and experimental evidence

Reverse transcriptase (RT) was first discovered as an essential catalyst in the biological cycle of retroviruses. However, in the past years evidence has accumulated showing that RTs are involved in a surprisingly large number of RNA-mediated transpositional events that include both viral and nonviral genetic entities. Although it is probable that some RT-bearing genetic elements like the different types of AIDS viruses and the mammalian LINE family have arisen in recent geological times, the possibility that reverse transcription first took place in the early Archean is supported by (1) the hypothesis that RNA preceded DNA as cellular genetic material; (2) the existence of homologous regions of the subunit tau of the E. coli DNA polymerase III with the simian immunodeficiency virus RT, the hepatitis B virus RT, and the beta' subunit of the E. coli RNA polymerase (McHenry et al. 1988); (3) the presence of several conserved motifs, including a 14-amino-acid segment that consists of an Asp-Asp pair flanked by hydrophobic amino acids, which are found in all RTs and in most cellular and viral RNA polymerases. However, whether extant RTs descend from the primitive polymerase involved in the RNA-to-DNA transition remains unproven. Substrate specificity of the AMV and HIV-1 RTs can be modified in the presence of Mn2+, a cation which allows them to add ribonucleotides to an oligo (dG) primer in a template-dependent reaction. This change in specificity is comparable to that observed under similar conditions in other nucleic acid polymerases. This experimentally induced change in RT substrate specificity may explain previous observations on the misincorporation of ribonucleotides by the Maloney murine sarcoma virus RT in the minus and plus DNA of this retrovirus (Chen and Temin 1980). Our results also suggest that HIV-infected macrophages and T-cell cells may contain mixed polynucleotides containing both ribo- and deoxyribonucleotides. The evolutionary significance of these changes in substrate specificities of nucleic acid polymerases is also discussed.

Non-NASA Center↗