Eye evolution at high resolution: The neuron as a unit of homology

Based on differences in morphology, photoreceptor-type usage and lens composition it has been proposed that complex eyes have evolved independently many times. The remarkable observation that different eye types rely on a conserved network of genes (including Pax6/eyeless) for their formation has led to the revised proposal that disparate complex eye types have evolved from a shared and simpler prototype. Did this ancestral eye already contain the neural circuitry required for image processing? And what were the evolutionary events that led to the formation of complex visual systems, such as those found in vertebrates and insects? The recent identification of unexpected cell-type homologies between neurons in the vertebrate and Drosophila visual systems has led to two proposed models for the evolution of complex visual systems from a simple prototype. The first, as an extension of the finding that the neurons of the vertebrate retina share homologies with both insect (rhabdomeric) and vertebrate (ciliary) photoreceptor cell types, suggests that the vertebrate retina is a composite structure, made up of neurons that have evolved from two spatially separate ancestral photoreceptor populations. The second model, based largely on the conserved role for the Vsx homeobox genes in photoreceptor-target neuron development, suggests that the last common ancestor of vertebrates and flies already possessed a relatively sophisticated visual system that contained a mixture of rhabdomeric and ciliary photoreceptors as well as their first- and second-order target neurons. The vertebrate retina and fly visual system would have subsequently evolved by elaborating on this ancestral neural circuit. Here we present evidence for these two cell-type homology-based models and discuss their implications.     

The eyes of a patrolling butterfly: visual field and eye structure in the Orange Sulphur, Colias eurytheme (Lepidoptera, Pieridae)

Sensory information plays a critical role in determining an animal's behavior on both proximate and evolutionary timescales. Butterflies, like many other insects, use vision extensively over their lifetimes, and yet relatively little work has been published to date on their visual capabilities. We describe the visual system of a pierid butterfly, Colias eurytheme, with the ultimate goal of better understanding its role in shaping the behavior of this animal. We made several measurements: visual field dimensions, eye surface area, interommatidial angle (Deltaphi), facet diameter (D), and eye parameter (p). C. eurytheme had a large visual field and considerable regional variation in visual acuity, as inferred by Deltaphi and D. When compared to females, males had larger eye surface areas, smaller Deltaphi, and larger D in all regions except ventrally. Both sexes had proportionally large eye surface areas compared to other butterflies. Minimum p in males was small, indicating that some regions of their eyes may operate close to the diffraction limit. Finally, we found that both eye surface area and D scaled positively, but with negative allometry to body size. We discuss the relevance of these visual characteristics to the biology and behavior of C. eurytheme.     

Genetic differentiation in life history between Daphnia galeata populations: an adaptation to local predation regimes?

Species of the water flea Daphnia exhibit constitutive as wellas phenotypically inducible anti-predator defence strategies,involving life history, morphological and behavioural traits.We explored the hypothesis of genetic differentiation in anti-predatordefence strategies using Daphnia clones originating from twodifferent water bodies: Tjeukemeer (the Netherlands) and FishPond (Belgium). Both water bodies are inhabited by zooplanktivorousfish. In contrast to Tjeukemeer, Fish Pond is also inhabitedby larvae of the phantom midge Chaoborus. The life history responsesof the two sets of clones to kairomones from fish (Perca), tokairomones from Chaoborus, and to a mixture of both were compared.Clones from Tjeukemeer and Fish Pond showed strong responsesto the presence of fish kairomone, with reductions in adultand neonate body length, in age at first reproduction and inthe total number of neonates produced during the first threeadult instars. Responses to Chaoborus kairomone were much lesspronounced, although there was a tendency towards an increasein the number of neonates in the first brood. Significant inter-populationgenetic differences were found for all the investigated traits.However, there was no indication for genetic adaptation of theFish Pond clones to negative size-selective predation by Chaoborus.Compared to Tjeukemeer clones, Fish Pond clones had a lowersize at first reproduction, produced smaller neonates and produceda higher number of neonates in their first brood. This suggestsadaptation to positive rather than to negative size selectivepredation. Genetic differences between populations were observedmainly for constitutive life history traits, rather than forphenotypic shifts in response to the presence of predator kairomones.     

Conjugation in Hyalophysa chattoni Bradbury (Apostomatida): An adaptation to a symbiotic life cycle

Hyalophysa chattoni, borne as an encysted phoront on a crustacean's exoskeleton, metamorphoses to the trophont during the host's premolt. After the molt within 15 min to 2 h conjugants with food vacuoles appear in the exuvium, swimming along with the trophonts. Starvation in other ciliates usually precedes conjugation, but food vacuoles in conjugants do not preclude starvation. Only after ingestion and dehydration of vacuoles ceases, does digestion of exuvial fluid begin. Conjugants resorb their feeding apparatus as they fuse. A single imperforate membrane from each partner forms the junction membrane. In a reproductive cyst conjugants divide synchronously, but now the junction membrane is interrupted by pores and channels. After the last division the daughters undergo meiosis – two meiotic divisions and one mitotic division yielding two prokarya as they simultaneously differentiate into tomites. After fertilization, pairs separate and the synkaryon divides once into a macronuclear anlage and a micronucleus. Exconjugants leave the cyst and seek a host. The parental macronucleus remains active until the phoront stage when the anlage develops. Owing to random association of micronuclei during meiosis, Hyalophysa's exconjugants are more genetically diverse than exconjugants from conventional patterns of conjugation.     

Higher vitellogenin concentrations in honey bee workers may be an adaptation to life in temperate climates

The honey bee originated in tropical Africa and later dispersed to northern Europe. It has been suggested that a higher hemolymph storage capacity for the glycolipoprotein vitellogenin evolved in temperate regions, and that the trait constitutes an adaptation to a strongly seasonal environment. We have investigated whether the relative vitellogenin levels of European and African honey bees are in accordance with this hypothesis. Our data indicate that European workers have a higher set-point concentration for vitellogenin compared to their African origin. Considered together with available life history information and physiological data, the results lend support to the view that “winter bees”, a longlived honey bee worker caste that survives winter in temperate regions, evolved through an increase in the worker bees’ capacity for vitellogenin accumulation. Received 20 September 2004; revised 25 March 2005; accepted 13 April 2005.     

Lobula plate and ocellar interneurons converge onto a cluster of descending neurons leading to neck and leg motor neuropil in Calliphora erythrocephala

Summary In the fly, Calliphora erythrocephala, a cluster of three Y-shaped descending neurons (DNOVS 1–3) receives ocellar interneuron and vertical cell (VS4–9) terminals. Synaptic connections to one of them (DNOVS 1) are described. In addition, three types of small lobula plate vertical cell (sVS) and one type of contralateral horizontal neuron (Hc) terminate at DNOVS 1, as do two forms of ascending neurons derived from thoracic ganglia. A contralateral neuron, with terminals in the opposite lobula plate, arises at the DNOVS cluster and is thought to provide heterolateral interaction between the VS4–9 output of one side to the VS4–9 dendrites of the other. DNOVS 2 and 3 extend through pro-, meso-, and metathoracic ganglia, branching ipsilaterally within their tract and into the inner margin of leg motor neuropil of each ganglion. DNOVS 1 terminates as a stubby ending in the dorsal prothoracic ganglion onto the main dendritic trunks of neck muscle motor neurons. Convergence of VS and ocellar interneurons to DNOVS 1 comprises a second pathway from the visual system to the neck motor, the other being carried by motor neurons arising in the brain. Their significance for saccadic head movement and the stabilization of the retinal image is discussed.     

Cyclomorphosis in Daphnia: an adaptation to avoid invertebrate predation

Seasonal morphological changes in three Daphnia species were followed over a two-year period in two lakes that differ in invertebrate and fish pressure. Whereas the morphology of D. hyalina, the biggest of the three species, varied little from season to season, D. cucullata, the smallest, exhibited the most pronounced seasonal changes in head height/carapace length ratio. The pattern of seasonal changes of body proportions was similar in all size classes and isometric growth of the head was reported for D. cucullata. Unlike the head, tail spine length/carapace length ratio almost did not vary seasonally. Strong negative allometry of tail spine growth was observed. These results are consistent with the hypothesis that helmets and tail spines provide protection against invertebrates in the two smallest, thus most endangered species.     

SMS 2.0: an updated database to study the structural plasticity of short peptide fragments in non-redundant proteins

The function of a protein molecule is greatly influenced by its three-dimensional (3D) structure and therefore structure prediction will help identify its biological function. We have updated Sequence, Motif and Structure (SMS), the database of structurally rigid peptide fragments, by combining amino acid sequences and the corre-sponding 3D atomic coordinates of non-redundant (25%) and redundant (90%) protein chains available in the Protein Data Bank (PDB). SMS 2.0 provides information pertaining to the peptide fragments of length 5-14 resi-dues. The entire dataset is divided into three categories, namely, same sequence motifs having similar, intermedi-ate or dissimilar 3D structures. Further, options are provided to facilitate structural superposition using the pro-gram structural alignment of multiple proteins (STAMP) and the popular JAVA plug-in (Jmol) is deployed for visualization. In addition, functionalities are provided to search for the occurrences of the sequence motifs in other structural and sequence databases like PDB, Genome Database (GDB), Protein Information Resource (PIR) and Swiss-Prot. The updated database along with the search engine is available over the World Wide Web through the following URL http://cluster.physics.iisc.ernet.in/sms/.     

Developmental instability as an index of adaptation to drought stress in a Mediterranean oak

An increase in temperature and water deficits caused by the ongoing climate change might lead to a decline growth rates and threaten the persistence of tree species in drought-prone areas within the Mediterranean Basin. Developmental instability (the error in development caused by stress) may provide an index of the adaptability of woody plants to withstand climatic stressors such as water shortage. This study evaluated the effects of drought stress on growth variables in three stands of a Mediterranean oak (Quercus faginea) exposed to differing climatic conditions (xeric, mesic and cooler) along an altitudinal gradient in northeastern Spain, in two climatically contrasting years (wet and dry years). Two indices of developmental instability, fluctuating and translational asymmetries, which reflect environmental stress, were measured in leaves and current-year shoots, respectively. We also measured branch biomass and fractal complexity of branches as indicators of the species’ performance. After a period of drought the individuals’ at the most xeric site presented lower developmental instability and less branch biomass than did the individuals from the mesic and cooler sites. We interpret that difference as an adaptive response to drought which reflects a trade-off between maintenance of homeostasis and growth when water is scarce. The study demonstrated that developmental instability constitutes a useful index to assess the degree of adaptation to stressful environmental conditions. The assessment of developmental instability in sites and years with contrasting climatic conditions provides a means of quantifying the capacity of plants to develop plastic adaptive responses to climatic stress.     

Skeletal economy in certain herbivorous beetles as an adaptation to a poor dietary supply of nitrogen

ABSTRACT.

• 1 Flying Coleoptera were sampled with light traps from the tree canopy of lowland alluvial rain forest in Brunei, Borneo.
• 2 Adult chrysomelid beetles were found to carry significantly less skeleton relative to their overall mass than those of any family other than herbivorous scarabaeids (subfamily: Rutelinae). Xylophagous lucanids and cerambycids and dung-feeding scarabaeids (subfamilies: Aphodiinae and Scarabaeinae) carried as large a proportion of skeleton by mass as predatory carabids and cicindelids.
• 3 Skeletal production requires the commitment of assimilated nitrogen to chitin and proteins. Adequate nitrogen is more readily available to predators, carrion or dung-feeders, and xylophages (given enough time) than to leaf-chewing chrysomelids, which are also environmentally exposed to predatory attack. Female chrysomelids were found to carry a larger number of mature eggs/insect than those of any other family. A larger proportion of their mass was represented by eggs than skeleton, as reproductive compensation for reduced skeletal protection.
• 4 It was estimated that a chrysomelid beetle required approximately 43% of the nitrogen incorporated by a predatory carabid of the same mass, whereas xylophagous cerambycids and lucanids would need between 84% and 98% of that in the predator.

     

Individuation in Sorghum bicolor: a self-organized process involved in physiological adaptation to salinity

In Sorghum bicolor, salt-adaptation was defined as the capacity of plants to grow at 300 m M NaCl (a lethal concentration) following a 3-week pre-treatment with 150 m M NaCl (a sublethal concentration). Large populations induced for salt-adaptation were analysed. Two modes of osmotic regulation (Na-includer and Na-excluder) were differentiated during induction of salt-adaptation, in spite of the genetic and environmental homogeneity. This variability was not related to any incapacity of the Na-includers to control accumulation of Na ⁺ ions in the shoot, because most of the pre-treated plants displayed a similar capacity of growth and control of Na ⁺ uptake after transfer to 300 m M NaCl. This unsuspected complexity of initially homogeneous populations points on the inadequacy of physiological studies focused on the ‘average individual’. Further analyses showed that variability was not directly related to micro-environmental variations. It is concluded that a process of individuation is caused by a third source of variability, which is the expression of a self-organizing process normally occurring during the transition phases in development. In constraining environments, this phenomenon of individuation includes adaptive modifications.     

Spectral broadening of the Soret band in myoglobin: an interpretation by the full spectrum of low-frequency modes from a normal modes analysis

In this work the temperature dependence of the Soret band line shape in carbon-monoxy myoglobin is re-analyzed by using both the full correlator approach in the time domain and the frequency domain approach. The new analyses exploit the full density of vibrational states of carbon-monoxy myoglobin available from normal modes analysis, and avoid the artificial division of the entire set of vibrational modes coupled to the Soret transition into "high-frequency" and "low-frequency" subsets; the frequency domain analysis, however, makes use of the so-called short-times approximation, while the time domain one avoids it. Time domain and frequency domain analyses give very similar results, thus supporting the applicability of the short-times approximation to the analysis of hemeprotein spectra; in particular, they clearly indicate the presence of spectral heterogeneity in the Soret band of carbon-monoxy myoglobin. The analyses also show that a temperature dependence of the Gaussian width parameter steeper than the hyperbolic cotangent law predicted by the Einstein harmonic oscillator and/or a temperature dependence of inhomogeneous broadening are not sufficient to obtain quantitative information on the magnitude of an-harmonic contributions to the iron-heme plane motion. However, the dependence of the previous two quantities may be used to obtain semiquantitative information on the overall coupling of the Soret transition to the low-frequency modes and therefore on the dynamic properties of the heme pocket in different states of the protein.     

Active sensing in a freely walking spider: look where to go

The Central American hunting spider Cupiennius salei, like most other spiders, has eight eyes, one pair of principal eyes and three pairs of secondary eyes. The principal eyes and one pair of the secondary eyes have almost completely overlapping visual fields, and presumably differ in function. The retinae of the principal eyes can be moved independently by two pairs of eye muscles each, whereas the secondary eyes do not have such eye muscles. The behavioural relevance of retinal movements of freely moving spiders was investigated by a novel dual-channel telemetric registration of the eye muscle activities. Walking spiders shifted the ipsilateral retina with respect to the walking direction before, during and after a turning movement. The change in the direction of vision in the ipsilateral anterior median eye was highly correlated with the walking direction, regardless of the actual light conditions. The contralateral retina remained in its resting position. This indicates that Cupiennius salei shifts it visual field in the walking direction not only during but sometimes previous to an intended turn, and therefore “peers” actively into the direction it wants to turn.     

How eukaryotic algae can adapt to the Spain's Rio Tinto: a neo-Darwinian proposal for rapid adaptation to an extremely hostile ecosystem

Microalgae contributed 60% of the total biomass in the extremely hostile (pH 2 and metal-rich waters) environment of Rio Tinto (which is used as a model for the astrobiology of Mars). These algae are closely related to nonextreme lineages, suggesting that adaptation to Rio Tinto water (RTW) must occur rapidly. Fitness from both the microalga Dictyosphaerium chlorelloides and the cyanobacterium Microcystis aeruginosa was inhibited when they were cultured in RTW. After further incubation for several weeks, D. chlorelloides survived, as a result of the growth of a variant that was resistant to RTW, but RTW-resistant cells did not appear in M. aeruginosa. A Luria-Delbrück fluctuation test revealed that D. chlorelloides RTW-resistant cells arose randomly by rare spontaneous mutations before the RTW exposure (1.38 x 10(-6) mutants per cell division). The mutants with a diminished fitness are maintained in nonextreme waters as the result of a balance between new RTW-resistant cells arising by mutation and RTW-resistant mutants eliminated by natural selection (equilibrium at c. 15 RTW-resistant per 10(7) wild-type cells). Rapid adaptation of eukaryotic algae to RTW could be the result of selection of RTW-resistant mutants occurring spontaneously in nonextreme populations that arrived fortuitously at the river in the past, or in the present continuously.     

Microbial adaptation to iron: a possible role of phosphatidylethanolamine in iron mineral deposition

Pseudomonas fluorescens multiplied in a minimal mineral medium supplemented with iron(III) (5 mm) complexed to citrate, the sole source of carbon, with no apparent diminution in cellular mass. Atomic absorption studies of different cellular fractions and supernatant at various growth intervals revealed that the trivalent metal was initially internalized. At approximately 41 h of incubation, the soluble cellular extract contained 9.5% of the iron originally found in the growth medium. However, as bacterial multiplication progressed, most of the metal was deposited as an extracellular insoluble gelatinous residue. Phosphatidylethanolamine appeared to be an important organic constituent of this precipitate. X-ray fluorescence and diffraction studies revealed that iron(III) was deposited as amorphous hydrated oxide. Scanning electron microscopy and energy dispersive X-ray microanalysis of the pellet aided in the identification of irregular shaped bodies rich in iron and oxygen that were associated with carbon-containing elongated structures. Examination of the bacterial cells by a transmission electron microscope equipped with an electron energy loss spectrometer indicated the deposition of iron within the cells.     

Testing for adaptation to climate in Arabidopsis thaliana: a calibrated common garden approach

BACKGROUND AND AIMS: A recent method used to test for local adaptation is a common garden experiment where analyses are calibrated to the environmental conditions of the garden. In this study the calibrated common garden approach is used to test for patterns of adaptation to climate in accessions of Arabidopsis thaliana. METHODS: Seedlings from 21 accessions of A. thaliana were planted outdoors in College Park, MD, USA, and development was monitored during the course of a growing season. ANOVA and multiple regression analysis were used to determine if development traits were significant predictors of plant success. Previously published data relating to accessional differences in genetic and physiological characters were also examined. Historical records of climate were used to evaluate whether properties of the site of origin of an accession affected the fitness of plants in a novel environment. KEY RESULTS: By calibrating the analysis to the climatic conditions of the common garden site, performance differences were detected among the accessions consistent with a pattern of adaptation to latitude and climatic conditions. Relatively higher accession fitness was predicted by a latitude and climatic history similar to that of College Park in April and May during the main growth period of this experiment. The climatic histories of the accessions were better predictors of performance than many of the life-history and growth measures taken during the experiment. CONCLUSIONS: It is concluded that the calibrated common garden experiment can detect local adaptation and guide subsequent reciprocal transplant experiments.     

Responses to environmental stressors in an estuarine fish: Interacting stressors and the impacts of local adaptation

Common killifish, Fundulus heteroclitus, are found in marshes and estuaries along the Atlantic coast of North America from Newfoundland to Florida. Although these habitats are highly productive, they are also characterized by variation in a number of abiotic stressors, including temperature, salinity, oxygen, and anthropogenic toxicants, which vary substantially in both space and time. In order to survive in these habitats, killifish must be able to cope with these stressors, both individually and in combination. There is substantial evidence to suggest that populations of F. heteroclitus have undergone local adaptation to multiple abiotic stressors, including temperature, salinity, and toxicants, but most studies have examined the effects of single stressors in isolation. Here I review some of the studies on local adaptation in F. heteroclitus, focusing on the molecular basis of local adaptation to abiotic stressors, and the acute responses to these stressors both singly and in combination. This work demonstrates that there are substantial interactions between the responses to both natural and anthropogenic stressors at the cellular level.     

Suppressed breeding in the field vole (Microtus agrestis): an adaptation to cyclically fluctuating predation risk

The densities of microtine rodents and their main predators,small mustelids, fluctuate synchronously in 3–5-year cyclesin central and northern Fennoscandia. Predation by small mustelidshas been suggested as one of the driving forces in microtinecyclicity, causing deep synchronous declines of several volespecies. We studied experimentally the effects of small mustelidson mating behavior, foraging, and breeding in nonwintered fieldvoles (Microtus agrestis) originating from a cyclic population.By using mustelid odors, we simulated a crash phase environmentwith high predation risk for breeding pairs of voles. In ourexperiments, 87% of the female field voles suppressed breedingwhen exposed to mustelid odors. Both female and male behaviorchanged, and no mating behaviors were observed under the simulatedpredation risk. Weights of both sexes decreased when exposedto mustelid odor, probably due to decreased foraging; weightsof the control females increased due to pregnancy; and no weightchanges occurred in control males. Decreased breeding and foragingpossibilities under high predation pressure may form the basisfor the ultimate explanation for breeding suppression. Thereare at least two different mechanisms for breeding suppression:either mating does not take place or malnutrition in femalesdoes not' allow breeding to occur. Delayed breeding under highrisk of predation, for whatever reason, could increase the probabilityof individuals, especially that of the females, to survive overthe crash to the next, safer breeding season when their youngwould have better possibilities to survive.     

Variation in the rate of convergent evolution: adaptation to a laboratory environment in Drosophila subobscura

Adaptation to novel environments is a central issue in evolutionary biology. One important question is the prevalence of convergence when different populations adapt to the same or similar environments. We investigated this by comparing two studies, 6 years apart, of laboratory adaptation of populations of Drosophila subobscura founded from the same natural location. In both studies several life‐history traits were periodically assayed for the first 14 generations of laboratory adaptation, as well as later generations, and compared with established, laboratory, control populations. The results indicated: (1) a process of convergence for all traits; (2) differences between the two studies in the pattern and rate of convergence; (3) dependence of the evolutionary rates on initial differentiation. The differences between studies might be the result of the differences in the founder populations and/or changes in the lab environment. In either case, the results suggest that microevolution is highly sensitive to genetic and environmental conditions.     

Microgeographic life history variation in an alpine caddisfly: plasticity in response to seasonal time constraints

1. Temporally constrained environments, such as habitats with short growth seasons or short hydroperiods, cause potentially strong selection on life histories. Depending on the predictability of these events and the extent of spatial and temporal heterogeneity, local populations could become adapted either via a fixed phenotype or via life history plasticity in response to these environmental cues. 2. We used a common garden experiment to investigate microgeographic variation in life history responses to combined changes in photoperiod (ambient/late) and hydroperiod (constant/drying) time constraint cues in an alpine caddisfly (Trichoptera). We compared six populations (three permanent/three temporary streams) originating from a small, alpine floodplain and which spanned an expected gradient in growth period duration (GPD) with distance from glaciers. 3. We made two main predictions in relation to locally varying selection pressures: (i) populations nearest glaciers (shorter GPD and strongest time constraints) should have the fastest development rates and (ii) populations from permanent streams should be less able to respond to drying hydroperiods than populations from temporary streams. 4. All populations and both sexes accelerated development in response to late season photoperiod cues. However, only permanent stream populations showed an increase in development time with increasing GPD, suggesting that other factors were influencing populations in temporary streams. 5. Permanent stream populations showed countergradient variation (genetic and environmental influences were in opposition) in development time, and under‐compensation of growth rates resulted in a converse Bergmann cline in body size (smaller body size along gradients of declining season length). The extent of plasticity in response to hydroperiod, and the combined effects of both time constraints, differed between populations and sexes, but were not consistent among populations. 6. Taken together, our results suggest adaptive plasticity in response to season length. The lack of a predictable pattern in response to hydroperiod may be due to gene flow or weak selection. We conclude that spatially structured populations can strongly differ in phenotypic plasticity even at microgeographic scales.