Evolution als eskalierende Individualisierung von Organismus und Umwelt

“Denn das Maß der Widerwärtigkeiten und Schlechtigkeiten wird augenblicklich wieder durch neue aufgefüllt, als glitte das eine Bein der Welt immer wieder zurück, wenn sich das andere vorschiebt. Daran müßte man die Ursache und den Geheimmechanismus erkennen!” (Robert Musil, Der Mann ohne Eigenschaften, I, 27) Die Frage, ob Evolution eine Fortschrittskomponente beinhalte, oder zumindest einen Trend zur Komplexität der Organismen oder der Ökosysteme, beschäftigt nicht nur Romanschreiber, sondern Biologen wie Philosophen (Ruse 1993). Ist Anpassung ein Schritt vorwärts? Wovon weg - worauf zu? Ist der komplexere Organismus besser an seine Umwelt angepaßt? Wenn ja, was sind die Konsequenzen und Kosten solchen Fortschritts? Ludwig von Bertalanffy wunderte sich über den Sinn eines evolutionären Dramas, in dem das Leben sich umständlich immer höher schraubt, um für jede erreichte Ebene einen neuen Preis zu zahlen: für Vielzelligkeit den Tod des Individuums, für das Nervensystem den Schmerz, für das Bewußtsein die Angst (Davidson 1983). Daß Evolution nicht, wie man lange glauben wollte, partout der Komplexität zustrebt, zeigt sich am Verlust der Flugfähigkeit bei vielen Inselvögeln in Abwesenheit terrestrischer Räuber, oder an der strukturellen Vereinfachung der meisten Parasiten, wird aber besonders deutlich in den klassischen Experimenten von Spiegelman (1967). Spiegelman inkubierte die RNA eines Virus in einer konstant gehaltenen Brühe aus freien Monomeren und Replikase. Unter artifizieller Selektion für rapide Reproduktion etablierte sich in diesem Experiment nach nur 75 Generationen eine stabile Mutante, die sich zwar fünfzehnmal so schnell vermehrte, sich aber von ursprünglich 4200 Nukleotiden auf nur mehr 220 reduziert hatte, nicht viel mehr als die Erkennungsstelle für die Replikase. Im Schlaraffenland, wo Ressourcen nie weniger werden, Abfälle sich nie anhäufen und Feinde nicht existieren, in einer Umwelt also, in welcher der Organismus weder bedroht noch von den Rückwirkungen seiner eigenen Handlungen betroffen wird, verläuft die Evolution anscheinend nicht in Richtung auf zunehmende Größe und Komplexität, wie dies Bonner (1988) für die reale Welt zu zeigen versuchte, sondern genau umgekehrt. Ich werde im folgenden eine Antwort auf dieses Paradoxon skizzieren, die aus den folgenden Thesen besteht: 1. Jeder Adaptivschritt eines evoluierenden Organismus zieht einen Komplementärschritt seiner unbelebten und seiner belebten Umwelt nach sich. Die Umwelt weicht vom sich adaptierenden Organismus zurück. Der Organismus von heute ist an die Umwelt von gestern angepaßt. 2. Während die unbelebte Umwelt lediglich in passiver Weise auf das energetische Vordringen des Organismus reagiert, sind Wechselwirkungen zwischen Organismus und belebter Umwelt, also anderen Organismen, teleonomisch. Wechselwirkungen, die im Energiefluß asymmetrisch sind (wie zum Beispiel zwischen Räuber und Beute), sind im Informationsfluß komplementär asymmetrisch. In dem Maße wie Energie zum erfolgreichen Räuber fließt, fließt Information zur erfolgreichen Beute. Der Räuber von heute erbt das Beutebild von gestern. 3. Einmalige Individualität, das Resultat sexueller Fortpflanzung, vermittelt diese asymmetrische Rückkoppelung und puffert die daraus resultierende zeitverschobene Eskalation. Der Trend zur Individualisierung ist selbstverstärkend und führt zu zunehmender räumlicher und zeitlicher Komplexität von Organismen und Umwelten. Individualität ist die stärkste Triebfeder in der Gestaltung der Biosphäre

Prometheus and Proteus: the creative, unpredictable individual in evolution

Evolutionary theory usually neglects two variables: the changes induced in the environment by the evolving organism, and individual uniqueness in sexually reproducing species. In order to fuel its maintenance and reproduction, an organism must average a positive net energy balance vis-a-v}s its environment. It achieves this via aptations, which consist of information (i.e., the internalization of all that is predictable about the environment, including the machinery to take advantage of this information) and stored energy (to operate the machinery, including a safety margin to deal with events that are unpredictable in principle). Taking advantage of a prediction, however, interferes with what has been predicted; each adaptation by the organism therefore changes its environmental target. Today's organism is adapted to yesterday's environment, and today's predator inherits yesterday's prey image. This paper attempts to show that, over evolutionary time, the persistence of this asymmetric, time-lagged relationship is owed increasingly to genetically unique individuals. Individual uniqueness as resulting from sexual reproduction is janusfaced. It endows an evolving population with both a forward-looking (promethean) and backward-looking (protean) feature. A population made up of genetically unique individuals is promethean (creative) in its ability to exploit non-homogeneous resources and respond serendipitously to environmental change via new genotypes; it is protean (elusive) in presenting a pursuer (predator or parasite) with a scattered target. Furthermore, because of the asymmetry between the winnowing of the target gene pool by the pursuer, and the genetic fixation in the pursuer of an outdated target image, the target keeps evolving away from the pursuer at a speed and in a direction that are a function of the pursuer's success. This mechanism ensures an evolutionary time lag be102 W. Sterrer tween pursuer and target, which explains escalation, the stability of asymmetric coevolutionary systems such as the life/dinner principle, and the pervasiveness of the Red Queen effect. Individuality thus both promotes and retards the speed of evolution. Having probably originated simultaneously with predation, sex-generated individuality is a self-accelerating evolutionary process that may account for much of today's organismic and environmental complexity

Detailed reconstruction of the nervous and muscular system of Lobatocerebridae with an evaluation of its annelid affinity

BACKGROUND: The microscopic worm group Lobatocerebridae has been regarded a ‘problematicum’, with the systematic relationship being highly debated until a recent phylogenomic study placed them within annelids (Curr Biol 25: 2000-2006, 2015). To date, a morphological comparison with other spiralian taxa lacks detailed information on the nervous and muscular system, which is here presented for Lobatocerebrum riegeri n. sp. based on immunohistochemistry and confocal laser scanning microscopy, supported by TEM and live observations. RESULTS: The musculature is organized as a grid of longitudinal muscles and transverse muscular ring complexes in the trunk. The rostrum is supplied by longitudinal muscles and only a few transverse muscles. The intraepidermal central nervous system consists of a big, multi-lobed brain, nine major nerve bundles extending anteriorly into the rostrum and two lateral and one median cord extending posteriorly to the anus, connected by five commissures. The glandular epidermis has at least three types of mucus secreting glands and one type of adhesive unicellular glands. CONCLUSIONS: No exclusive “annelid characters” could be found in the neuromuscular system of Lobatocerebridae, except for perhaps the mid-ventral nerve. However, none of the observed structures disputes its position within this group. The neuromuscular and glandular system of L. riegeri n. sp. shows similarities to those of meiofaunal annelids such as Dinophilidae and Protodrilidae, yet likewise to Gnathostomulida and catenulid Platyhelminthes, all living in the restrictive interstitial environment among sand grains. It therefore suggests an extreme evolutionary plasticity of annelid nervous and muscular architecture, previously regarded as highly conservative organ systems throughout metazoan evolution. ELECTRONIC SUPPLEMENTARY MATERIAL: The online version of this article (doi:10.1186/s12862-015-0531-x) contains supplementary material, which is available to authorized users

An Anatomical Description of a Miniaturized Acorn Worm (Hemichordata, Enteropneusta) with Asexual Reproduction by Paratomy

The interstitial environment of marine sandy bottoms is a nutrient-rich, sheltered habitat whilst at the same time also often a turbulent, space-limited, and ecologically challenging environment dominated by meiofauna. The interstitial fauna is one of the most diverse on earth and accommodates miniaturized representatives from many macrofaunal groups as well as several exclusively meiofaunal phyla. The colonization process of this environment, with the restrictions imposed by limited space and low Reynolds numbers, has selected for great morphological and behavioral changes as well as new life history strategies. Here we describe a new enteropneust species inhabiting the interstices among sand grains in shallow tropical waters of the West Atlantic. With a maximum body length of 0.6 mm, it is the first microscopic adult enteropneust known, a group otherwise ranging from 2 cm to 250 cm in adult size. Asexual reproduction by paratomy has been observed in this new species, a reproductive mode not previously reported among enteropneusts. Morphologically, Meioglossus psammophilus gen. et sp. nov. shows closest resemblance to an early juvenile stage of the acorn worm family Harrimaniidae, a result congruent with its phylogenetic placement based on molecular data. Its position, clearly nested within the larger macrofaunal hemichordates, suggests that this represents an extreme case of miniaturization. The evolutionary pathway to this simple or juvenile appearance, as chiefly demonstrated by its small size, dense ciliation, and single pair of gill pores, may be explained by progenesis. The finding of M. psammophilus gen. et sp. nov. underscores the notion that meiofauna may constitute a rich source of undiscovered metazoan diversity, possibly disguised as juveniles of other species.Organismic and Evolutionary Biolog

Microanatomy of the trophosome region of Paracatenula cf. polyhymnia (Catenulida, Platyhelminthes) and its intracellular symbionts

Marine catenulid platyhelminths of the genus Paracatenula lack mouth, pharynx and gut. They live in a symbiosis with intracellular bacteria which are restricted to the body region posterior to the brain. The symbiont-housing cells (bacteriocytes) collectively form the trophosome tissue, which functionally replaces the digestive tract. It constitutes the largest part of the body and is the most important synapomorphy of this group. While some other features of the Paracatenula anatomy have already been analyzed, an in-depth analysis of the trophosome region was missing. Here, we identify and characterize the composition of the trophosome and its surrounding tissue by analyzing series of ultra-thin cross-sections of the species Paracatenula cf. polyhymnia. For the first time, a protonephridium is detected in a Paracatenula species, but it is morphologically reduced and most likely not functional. Cells containing needle-like inclusions in the reference species Paracatenula polyhymnia Sterrer and Rieger, 1974 were thought to be sperm, and the inclusions interpreted as the sperm nucleus. Our analysis of similar cells and their inclusions by EDX and Raman microspectroscopy documents an inorganic spicule consisting of a unique magnesium–phosphate compound. Furthermore, we identify the neoblast stem cells located underneath the epidermis. Except for the modifications due to the symbiotic lifestyle and the enigmatic spicule cells, the organization of Paracatenula cf. polyhymnia conforms to that of the Catenulida in all studied aspects. Therefore, this species represents an excellent model system for further studies of host adaptation to an obligate symbiotic lifestyle

The Magnitude of Global Marine Species Diversity

Background: The question of how many marine species exist is important because it provides a metric for how much we do and do not know about life in the oceans. We have compiled the first register of the marine species of the world and used this baseline to estimate how many more species, partitioned among all major eukaryotic groups, may be discovered. Results: There are ∼226,000 eukaryotic marine species described. More species were described in the past decade (∼20,000) than in any previous one. The number of authors describing new species has been increasing at a faster rate than the number of new species described in the past six decades. We report that there are ∼170,000 synonyms, that 58,000–72,000 species are collected but not yet described, and that 482,000–741,000 more species have yet to be sampled. Molecular methods may add tens of thousands of cryptic species. Thus, there may be 0.7–1.0 million marine species. Past rates of description of new species indicate there may be 0.5 ± 0.2 million marine species. On average 37% (median 31%) of species in over 100 recent field studies around the world might be new to science. Conclusions: Currently, between one-third and two-thirds of marine species may be undescribed, and previous estimates of there being well over one million marine species appear highly unlikely. More species than ever before are being described annually by an increasing number of authors. If the current trend continues, most species will be discovered this century

Lurus minos, the first species of Luridae (Turbellaria: Rhabdocoela) from the Old World

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Haplognathia asymmetrica Sterrer 1991

<i>Haplognathia asymmetrica</i> Sterrer, 1991 <p>(Figs.1.1–1.13, 2.1–2.3)</p> <p> <i>Material</i> 10 specimens (one adult, and 9 juveniles or anterior fragments) from samples SI 10 and SI 16.</p> <p> <i>Distribution</i> Hawaii (Sterrer 1991b), NE Australia (Sterrer 2001), and (sub)tropical NW Atlantic (Sterrer 1998).</p> <p> <i>Description</i></p> <p>Colorless­translucent, often with many refractile granula in the intestine. The only adult (Fig. 1.1) was 1710 µm long and 35 µm wide at U 23.4 (index 48.86), with a pointed rostrum 220 µm long and 35 µm wide at U 8.8 (index 6.29). A single mature egg, 185 µm long and 15 µm wide, extended from U 39 to U 50. This specimen contained a bundle of allosperm at about U 25. The paired testes begin at about U 60; paired vasa deferentia open into a glandular male pore situated ventrally, at U 95, 90 µm anterior to the thin tail end.</p> <p>The basal plate is asymmetric, triangular, 9.5 µm long and 5.7 µm wide (index 1.74). Its dorsal surface bears two longitudinal ridges. Its posterior corner is usually, and its rostrolateral corners are sometimes knob­shaped. Jaws are 22.20 µm long, toothless, compact, with short, horn­shaped rostral apophyses and a transversely oval symphysis. The jaws of one specimen showed grainy degeneration (Fig. 1.7). The pharynx measures 4 µm behind the symphysis.</p> <p> <i>Discussion</i></p> <p> Most similar to <i>H. simplex</i> (Sterrer, 1969), this species is easily identified by its asymmetric basal plate. New Zealand specimens are in agreement with records from other localities (jaw lengths: Hawaii 26.67 µm, Australia 22.67 µm, NW Atlantic 21.93 µm) except by having a considerably narrower basal plate, as expressed in the high basal plate index of 1.74 (Hawaii 1.42, Australia 1.17, NW Atlantic 1.21).</p>Published as part of <i>Sterrer, Wolfgang, 2006, Gnathostomulida from the Otago Peninsula, southern New Zealand, pp. 1-19 in Zootaxa 1172</i> on pages 3-5, DOI: <a href="http://zenodo.org/record/2645647">10.5281/zenodo.2645647</a&gt

Pterognathia ugera Sterrer 1991

<i>Pterognathia ugera</i> Sterrer, 1991 <p>(Fig.Fig. 4.4–4.6)</p> <p> <i>Material</i></p> <p>Four juveniles/anterior fragments from sample SI 16.</p> <p> <i>Distribution</i></p> <p>Tahiti (Sterrer 1991c), (sub)tropical NW Atlantic (Sterrer 1998).</p> <p> <i>Description</i></p> <p>Rostrum measurements of this colorless­opaque species were 175–270 µm in length and 45–50 µm in width (index 4.64). The pharynx is 12.67 µm long behind the symphysis. The basal plate, 7.75 µm long and 11.25 µm wide (index 0.70) is horseshoe­shaped, bearing a pair of latero­caudally pointing wings; its rostral rim is entirely set with 25–31 (28.00) very regular teeth. The jaws are compact, 13.75 µm long, with short rostral apophyses and 3–4 teeth. In unsqueezed specimens the basal plate covers the jaws such that the jaw apophyses coincide with the latero­caudal extensions of the basal plate (Fig. 4.5).</p> <p> <i>Discussion</i> Easily identified by its unique basal plate, this circumtropical species may eventually warrant a separate genus due to its distinctive mouth part architecture (Sterrer 1998).</p> <p> <b> <i>Pterognathia portobello</i> n. sp.</b> </p> <p>(Fig.4.7–4.8)</p> <p> <i>Type material</i> Holotype one anterior fragment from sample SI 16, in squeeze preparation, NZNM W.1535.</p> <p> <i>Etymology</i> In appreciation of facilities and help provided during my stay at the Portobello Marine Laboratory, University of Otago, Dunedin, New Zealand.</p> <p> <i>Diagnosis</i></p> <p> <i>Pterognathia</i> with delicate jaws 13 µm long. Basal plate 2 µm long and 14 µm wide (index 0.14), with several rows of many minute teeth over median two thirds of rostral edge.</p> <p> <i>Description</i></p> <p>The only specimen, an anterior fragment, was colorless­translucent, 1350 µm long and 50 µm wide, and had a rostrum 270 µm long and 35 µm wide (index 7.71).</p> <p>The basal plate is a thin transverse sliver, 2 µm long and 14 µm wide (index 0.14), with pointed lateral tips. The median two thirds of its rostral edge are set with two or more horizontal rows of minute thorn­like teeth. The jaws are 13 µm long and delicate. The narrow symphysis lamellae join posteriorly in a small, globular symphysis. Anteriorly the jaws broaden, but it was not possible to identify distinct features such as rostral apophyses or cristae. Each jaw terminates dorso­rostrally in a knob, and ventro­rostrally seemed to have 4–5 teeth. The pharynx measures 10 µm in length behind the symphysis.</p> <p>Nothing is known about the reproductive system.</p> <p> <i>Discussion</i></p> <p> A much­wider­than­long basal plate and many­toothed jaws characterize <i>Cosmognathia</i> and <i>Pterognathia</i>. In its ‘front­heavy’ jaw architecture the new species most resembles <i>Pterognathia ugera</i> Sterrer, 1991. Although the single specimen represents without doubt a new species, the fragmentary and delicate nature of its features suggests leaving the assignment to genus tentative pending an analysis of additional material.</p>Published as part of <i>Sterrer, Wolfgang, 2006, Gnathostomulida from the Otago Peninsula, southern New Zealand, pp. 1-19 in Zootaxa 1172</i> on pages 9-11, DOI: <a href="http://zenodo.org/record/2645647">10.5281/zenodo.2645647</a&gt

Haplognathia ruberrima

<i>Haplognathia ruberrima</i> (Sterrer, 1966) <p>(Fig.3.7–3.8)</p> <p> <i>Material</i> Three juveniles/anterior fragments from sample SI 16.</p> <p> <i>Distribution</i></p> <p>North Sea, Adriatic (Sterrer 1969), Canary Islands (Sterrer 1997), (sub)tropical NW Atlantic (Sterrer 1998), Fiji (Sterrer 1991a), Hawaii (Sterrer 1991b), NE Australia (Sterrer 2001).</p> <p> <i>Description</i></p> <p>Body bright crimson. Basal plate more or less hexagonal, 5.67 µm long, 8.00 µm wide (index 0.71), with rows of thorns on its dorsal surface. Jaws 20.00 µm long, with one pair of teeth and with relatively long, straight rostral apophyses (apophysis index 0.50).</p> <p> <i>Discussion</i></p> <p> Sterrer (1998) discussed the cosmopolitan distribution, variability, frequent sympatry, and possible tendency of <i>H. ruberrima</i> and <i>H. rosea</i> to hybridize. The two species are nevertheless fairly distinguished by jaw apophysis index (0.50 or more in <i>H. ruberrima</i>, 0.50 or less in <i>H. rosea</i>) and details of the basal plate (with dorsal thorns in <i>H. ruberrima</i>, without thorns but with longitudinal ridges in <i>H. rosea</i>).</p> <p> <b>Family Pterognathiidae Sterrer, 1972</b></p> <p> <b> <i>Pterognathia sica</i> Sterrer, 2001</b> </p> <p>(Fig.4.1–4.3)</p> <p> <i>Material</i> Two juveniles from sample SI 16.</p> <p> <i>Distribution</i> NE Australia (Sterrer 2001).</p> <p> <i>Description</i></p> <p>Colorless­translucent. The larger of the juveniles was 1150 µm long and 45 µm wide (index 25.5), with a rostrum 150 µm long and 30 µm wide (index 5.00). Neither of the two specimens had a basal plate. The jaws are slender, and 18 µm long in both specimens; with long, curved rostral apophyses, dorsal cristae, and a small, transverse­oval symphysis. There are 5 long, ventral teeth and 3–4 shorter dorsal teeth.</p> <p> <i>Discussion</i></p> <p> Although the jaws of the NZ specimens are considerably shorter than those of Australian specimens (26.00 µm), they otherwise agree well with the original description of <i>P. sica</i>, the only known species in the genus <i>Pterognathia</i> that consistently lacks a basal plate.</p>Published as part of <i>Sterrer, Wolfgang, 2006, Gnathostomulida from the Otago Peninsula, southern New Zealand, pp. 1-19 in Zootaxa 1172</i> on pages 7-9, DOI: <a href="http://zenodo.org/record/2645647">10.5281/zenodo.2645647</a&gt