A hexapedal jointed-leg model for insect locomotion in the horizontal plane

We develop a simple model for insect locomotion in the horizontal (ground) plane. As in earlier work by Seipel et al. (Biol Cybern 91(0):76–90, 2004) we employ six actuated legs that also contain passive springs, but the legs, with “hip” and ‘knee’ joints, better represent insect morphology. Actuation is provided via preferred angle inputs at each joint, corresponding to zero torques in the hip and knee springs. The inputs are determined from estimates of foot forces in the cockroach Blaberus discoidalis via an inverse problem. The head–thorax–body is modeled as a single rigid body, and leg masses, inertia and joint dissipation are ignored. The resulting three degree-of-freedom dynamical system, subject to feedforward joint inputs, exhibits stable periodic gaits that compare well with observations over the insect’s typical speed range. The model’s response to impulsive perturbations also matches that of freely-running cockroaches (Jindrich and Full, J Exp Biol 205:2803–2823, 2002), and stability is maintained in the face of random foot touchdowns representative of real insects. We believe that this model will allow incorporation of realistic muscle models driven by a central pattern generator in place of the joint actuators, and that it will ultimately permit the study of proprioceptive feedback pathways involving leg force and joint angle sensing.     

A temperature-sensitive DNA adenine methyltransferase mutant of Salmonella typhimurium

A temperature-sensitive mutant of Salmonella typhimurium was isolated earlier after transposon mutagenesis with Tn10d Tet. The mutant D220 grows well at 28 °C but has a lower growth rate and forms filaments at 37 °C. Transposon-flanking fragments of mutant D220 DNA were cloned and sequenced. The transposon was inserted in the dam gene between positions 803 and 804 (assigned allele number: dam-231 : : Tn10d Tet) and resulted in a predicted ten-amino-acid-shorter Dam protein. The insertion created a stop codon that led to a truncated Dam protein with a temperature-sensitive phenotype. The insertion dam-231 : : Tn10d Tet resulted in a dam“leaky” phenotype since methylated and unmethylated adenines in GATC sequences were present. In addition, the dam-231 : : Tn10d Tet insertion rendered dam mutants temperature-sensitive for growth depending upon the genetic background of the S. typhimurium strain. The wild-type dam gene of S. typhimurium exhibited 82% identity with the Escherichia coli dam gene.     

Binding and segmentation of multiple objects through neural oscillators inhibited by contour information

 Temporal correlation of neuronal activity has been suggested as a criterion for multiple object recognition. In this work, a two-dimensional network of simplified Wilson-Cowan oscillators is used to manage the binding and segmentation problem of a visual scene according to the connectedness Gestalt criterion. Binding is achieved via original coupling terms that link excitatory units to both excitatory and inhibitory units of adjacent neurons. These local coupling terms are time independent, i.e., they do not require Hebbian learning during the simulations. Segmentation is realized by a two-layer processing of the visual image. The first layer extracts all object contours from the image by means of “retinal cells” with an “on-center” receptive field. Information on contour is used to selectively inhibit Wilson-Cowan oscillators in the second layer, thus realizing a strong separation among neurons in different objects. Accidental synchronism between oscillations in different objects is prevented with the use of a global inhibitor, i.e., a global neuron that computes the overall activity in the Wilson-Cowan network and sends back an inhibitory signal. Simulations performed in a 50×50 neural grid with 21 different visual scenes (containing up to eight objects + background) with random initial conditions demonstrate that the network can correctly segment objects in almost 100% of cases using a single set of parameters, i.e., without the need to adjust parameters from one visual scene to the next. The network is robust with reference to dynamical noise superimposed on oscillatory neurons. Moreover, the network can segment both black objects on white background and vice versa and is able to deal with the problem of “fragmentation.” The main limitation of the network is its sensitivity to static noise superimposed on the objects. Overcoming this problem requires implementation of more robust mechanisms for contour enhancement in the first layer in agreement with mechanisms actually realized in the visual cortex. Received: 25 October 2001 / Accepted: 26 February 2003 / Published online: 20 May 2003 Correspondence to: Mauro Ursino (e-mail: mursino@deis.unibo.it, Tel.: +39-051-2093008, Fax: +39-051-2093073)     

Detection of QTLs for crossability in wheat using a doubled-haploid population

 An intervarietal molecular-marker map was used for the detection of genomic regions influencing crossability between wheat (Triticum aestivum L. em Thell) and rye (Secale cereale L.). Analysis of deviance and logistic marker-regression methods were conducted on data from doubled haploid lines from a cross between “Courtot” and “Chinese Spring”. A major quantitative trait locus (QTL) involved in crossability, associated with the marker Xfba367-5B, was detected on the short arm of chromosome 5B. An additional locus, Xwg583-5B, was indicated on the long arm of chromosome 5B. This minor QTL might correspond to Kr1 which was presumed to be the major gene controlling crossability. Another locus of the genome, Xtam51-7A on chromosome 7A, was significantly associated with this trait. Alleles of “non-crossability” were contributed by the non-crossable cultivar “Courtot”. The three-marker model explains 65% of the difference in crossability between the two parents. The present results are discussed in relation to those previously carried out to locate the Kr genes by using the telocentric mapping technique. Received: 27 February 1998 / Accepted: 15 May 1998     

Computational model of the cockroach escape behavior: winner and losers in a population code

 I present a comprehensive biologically oriented computational model to account for the escape response of the cockroach on the ground. This model is an expansion of previous work that accounted only for discriminating left from right wind directions [Ezrachi et al. (1999) Biol Cybern 81: 89–99]. The model is composed of computational elements describing the biological processes taking place in the various neurons and includes input which emulates empirical data. With this model it is possible to obtain escape behavior that resembles natural behavior. The model is used to address an ongoing debate as to whether the cockroach's turn direction is determined by computations carried out by the entire neuronal population (PC) or rather by a “winner-take-all” (WTA) mechanism. I suggest that the computation mechanism that underlies the cockroach escape response is composed of both PC and WTA principles. Based on the properties of the suggested new mechanism I denote it a “Darwinian population code.” Received: 26 March 2002 / Accepted in revised form: 24 June 2002 Acknowledgements. I thank H. Parnas for her advice and assistance, J. M. Camhi for helpful comments, and D. Lipson for developing the simulation tools. Correspondence to: E. A. Ezrachi (e-mail: erez@piano.ls.huji.ac.il, Tel.: +972-2-6585818, Fax: +972-2-6585569)     

“Bacillus thermoantarcticus” sp. nov., from Mount Melbourne,Antarctica: a novel thermophilic species

 A novel thermophilic Gram-positive bacillus, “Bacillus thermoantarcticus”, isolated from geothermal soil near the crater of Mount Melbourne, is described. The organism grows at an optimal temperature of 63^°C at pH 6.0, is oxidase-positive, catalase-negative and produces an exopolysaccharide, an exocellular xylanase, an intracellular alcohol dehydrogenase and exo- and endocellular α-glucosidase(s). The sequence of 16S rDNA is very similar to that of “Bacillus thermoglucosidasius”; however, the guanine-plus-cytosine (G+C) content is 8 mol% higher. The type strain is “Bacillus thermoantarcticus” (DSM 9572). Received : 3 February 1995/Accepted : 12 May 1995     

Salt tolerance in Lycopersicon species VI. Genotype-by-salinity interaction in quantitative trait loci detection: constitutive and response QTLs

 A study of genotype-by-salinity interaction was carried out to compare the behavior of quantitative trait loci (QTLs) in two F₂ populations derived from crosses between the cherry tomato, Lycopersicon esculentum Mill. var. cerasiforme, and two wild relatives Lycopersicon pimpinellifolium (Jusl.) Mill. and Lycopersicon chesmannii f. minor (Hook. f.) Mull., grown at two environmental conditions (optimum and high salinity). QTLs for earliness and fruit yield could be classified into four groups: “response-sensitive”, those detected only under control conditions or whose contribution significantly decreased in salinity; “response-tolerant”, detected only in salinity or in which the direction of their additive effects changed; “constitutive”, detected in both growing conditions; and “altered” QTLs, those where the degree of dominance changed according to the presence or absence of salt. Epistatic interactions were also influenced by the salt treatment. This differential allele effect at some (non-constitutive) QTLs induced by salt stress will make selection under an “optimum environment” unfruitful for the “response-tolerant” QTLs. Similarly, selection under salinity will ignore “response-sensitive” QTLs. Given that salinity is highly variable in the field, marker-assisted selection should take into account not only the “response-tolerant” but also the “response-sensitive” QTLs although there might be cases where selection in some QTLs for both conditions is not feasible. Comparing both populations, very few QTLs showed the same behavior. Received: 5 August 1996 / Accepted: 25 October 1996     

Continuous cultures of tomato and citron roots in vitro

Summary Initial trials with tomato-root cultures disclosed the desirability of employing a gently agitated liquid medium containing iron in the chelated form. For the normal cultivars “Ace” and “Tropic”, subcultures were best achieved by utilizing sectors that possessed one or more newly emerged laterals. Continuous cultures of a nonlateral-forming tomato mutant, “Diageotropica”, and of citron were accomplished by subculturing tips of the elongating primary roots. The tomato roots were cultured in White's medium with the Fe₂(SO₄)₃ replaced by 0.03 mM NaFeEDTA. Sustained growth of citron-root tips necessitated the use of a medium containing Murashige and Skoog salts, 7.5% sucrose, 100 mg per I each of citric acid and thiamine HCl, and 5000 mg per li-inositol. The success with citron-root cultures was extendable to all cultivars ofC. medica L., but not to otherCitrus species relatives. Both citron and “Diageotropica” root cultures manifested undiminished elongation through repeated subcultures; but neither produced laterals in response to any cultural treatments. Research was supported in part by National Science Foundation Grant OIP75-10390 and Elvenia J. Slosson Fellowship in Ornamental Horticulture.     

Synthesis and paralytic activities of squaryl amino acid-containing polyamine toxins

Summary.  Eight analogs 4a-7a and 4b-7b of philanthotoxin (PhTX) from wasp venom and nephilatoxin-8 (NPTX-8) from spider venom whose tyrosine or asparagine linker is replaced by squaryl (sq) amino acid or 4-amino squaryl (4-asq) amino acid have been synthesized in an efficient manner via coupling of N-acyl squaryl amino acid intermediate 19 or 26 with the corresponding polyamine part. Preliminary bioassay using crickets revealed that the analogs substituted by glutamate-type squaryl amino acid-containing NPTX 7a and 7b showed more potent paralytic activities than that of NPTX-8. Received April 25, 2002 Accepted June 21, 2002 Published online December 18, 2002 Acknowledgement This work was supported by a grant from Research for the Future Program from the Japan Society for the Promotion of Science (JSPS). Authors' address: Yasufumi Ohfune, Graduate School of Science, Osaka City University, Sugimoto, Osaka 558-8585, Japan, Fax: +81-6-6605-3153, E-mail: ohfune@sci.osaka-cu.ac.jp     

Information-theoretic analysis of de-efferented single muscle spindles

 The information transmission properties of single, de-efferented primary muscle-spindle afferents from the hind limb of the cat were investigated. The gastrocnemius medialis muscle was stretched randomly while recording spike trains from several muscle-spindle afferents in the dorsal root. Two classes of input stimuli were used: (i) Gaussian noise with band-limited flat spectrum, and (ii) Gaussian noise with a more “naturalistic” 1/f ⁿ spectrum. The “reconstruction” method was used to calculate a lower bound to the information rate (in bits per second) between the muscle spindles and the spinal cord. Results show that in response to the flat-spectrum input, primary muscle-spindle afferents transfer information mainly about high frequencies, carrying 2.12 bits/spike. In response to naturalistic-spectrum inputs, primary muscle-spindle afferents transfer information about both low and high frequencies, with “spiking efficiency” increasing to 2.67 bits/spike. A simple muscle-spindle simulation model was analyzed with the same method, emphasizing the important part played by the intrafusal fiber mechanical properties in information transmission. Received: 22 January 2002 / Accepted in revised form: 17 June 2002 Correspondence to: Y. Tock (e-mail: ytock@tx.technion.ac.il, Fax: +972-4-8323041)     

Does age matter in studying the crustacean eye?

 Fifty randomly selected publications dealing with aspects of crustacean vision had their Materials and methods sections examined. In half of the articles gleaned, only the name of the animal under study was given; the other half mentioned “adult” or “mature” animals; twelve papers provided information on size or age of the experimental animal and six gave the sex. In Petrolisthes elongatus, rhabdom microvilli not only become more regular in outline as the animal grows, but also decrease in diameter from 90.4±21.2 nm in crabs of 5.0–8.9 mm carapace length via 86.6±13 nm in crabs of 9.0–12.9 mm carapace length to 79.7±7 nm in crabs of 13.0–16.9 mm carapace length. Approximately 400 cross sectional diameters of microvilli from identical regions in four eyes of each size class were measured and provided the basis for Anova-, Levene-, and t-tests. The three size classes possess microvilli of significantly different diameters and standard deviations. Our observations show that investigators of the crustacean photoreceptor have frequently neglected to consider the size of their experimental animals, but that this practice should not be continued as the now documented changes in microvillus diameters may influence the amount of visual pigment present and, thus, the photoreceptor’s sensitivity to light. Accepted: 7 May 1996     

Genetic control of aluminium tolerance in rye (Secale cereale L.)

 Aluminium (Al) tolerance in roots of two cultivars (“Ailés” and “JNK”) and two inbred lines (“Riodeva” and “Pool”) of rye was studied using intact roots immersed in a nutrient solution at a controlled pH and temperature. Both the cultivars and the inbred lines analysed showed high Al tolerance, this character being under multigenic control. The inbred line “Riodeva” was sensitive (non-telerant) at a concentration of 150 μM, whereas the “Ailes” cultivar showed the highest level of Al tolerance at this concentration. The segregation of aluminium-tolerance genes and several isozyme loci in different F₁s, F₂s and backcrosses between plants of “Ailés” and “Riodeva” were also studied. The segregation ratios obtained for aluminium tolerance in the F₂s analysed were 3 : 1 and 15 : 1 (tolerant : non-tolerant) while in backcrosses they were 1 : 1 and 3 : 1. These results indicated that Al tolerance is controlled by, at least, two major dominant and independent loci in rye (Alt1 and Alt3). Linkage analyses carried out between Al-tolerance genes and several isozyme loci revealed that the Alt1 locus was linked to the aconitase-1 (Aco1), nicotinamide adenine dinucleotide dehydrogenase-2 (Ndh2), esterase-6 (Est6) and esterase-8 (Est8) loci, located on chromosome arm 6RL. The order obtained was Alt1-Aco1-Ndh2-Est6-Est8. The Alt3 locus was not linked to the Lap1, Aco1 and Ndh2 loci, located on chromosome arms, 6RS, 6RL and 6RL respectively. Therefore, the Alt3 locus is probably on a different chromosome. Received: 18 March 1997 / Accepted: 21 March 1997     

Implementation of a small-scale “no-use zone” policy in a reef ecosystem: Eilat’s reef-lagoon six years later

 A small-scale, “no-use zone policy” has been implemented since 1992 at Eilat’s Coral Nature Reserve (Northern Red Sea). Six years later, the status of this closed-to-the-public reef area was compared to two nearby open-to-the-public sites, by evaluating populations of the scleractinian coral Stylophora pistillata in the strolling zone (0.5–1.5 m depth). Results from the open sites show that: (1) Live coral cover was three times lower than at the closed site; (2) numbers of small colonies (recruits) were significantly higher than in the closed site, while numbers of medium and large size colonies (geometric mean radius, rˉ>4.1 cm) per m² were significantly lower; (3) maximum rˉ was almost half than that in the closed site (9.6 cm versus 16.7 cm); (4) average number of broken colonies was three times higher than in the closed site; (5) significantly fewer colonies were partially dead. The latter result may reflect senescence processes in the large colonies of the closed site. Although colony breakage is reduced, it appears that the “no-use zone” policy is not sufficient for protecting small reef areas. The intense exploitation of Eilat’s coral reef by the tourist industry requires’ in addition to the conventional protective measures, the initiation of novel management solutions such as reef restoration by sexual and asexual recruits. Accepted: 11 August 1999     

3-Carboxy-4-phosphonocyclopentane amino acids: New metabotropic glutamate receptor ligands

Summary.  Two glutamic acid analogs (1SR,3RS,4RS)- and (1SR,3SR,4SR)-1-amino-4-phosphono cyclopentane-1,3-dicarboxylic acids (APCPD) have been synthesized. Pure E-(diethoxy-phosphoryl)-acrylic acid ethyl ester was obtained from ethyl propiolate, phenol and triethylphosphite. It was used as dienophile in a Diels-Alder reaction. Oxidation and cyclization afforded 3-(ethoxy-carbonyl)-4-(diethoxy-phosphoryl)-cyclopentanone. Bucherer-Bergs reaction and hydrolysis yielded APCPD-III and -IV which are inactive on mGlu1a receptor and antagonists on mGlu2 and mGlu8a receptors. Received April 2, 2002 Accepted July 11, 2002 Published online December 18, 2002 Acknowledgments This work was supported by grants from the CNRS, the programs “Physique et Chimie du Vivant” (PCV00–134, CNRS) and “Molécules et Cibles Thérapeutiques” (CNRS/INSERM), RETINA France and the Fondation de France (Comité Parkinson). A.-S. B. was supported by fundings from Parke-Davis Pharmaceutical Research (Ann Arbor, MI) and the Fondation de la Recherche Médicale. Authors' address: Dr Francine C. Acher, Laboratoire de Chimie et Biochimie Pharmacologiques et Toxicologiques, UMR8601-CNRS, Université René Descartes-Paris V, 45 rue des Saints-Pères, 75270 Paris Cedex 06, France, Fax: (33) 1 42 86 83 87, E-mail: acher@biomedicale.univ-paris5.fr     

A space-variant filter model of texture segregation: Parameter adjustment guided by psychophysical data

 This article presents a space-variant version of a standard spatial filter model of texture segregation of the “back-pocket” type (i.e., two filter layers with an intermediate pointwise nonlinearity). The model was tested with psychophysical data from experiments with line textures in which target lines differed in orientation from background lines. The textures were presented briefly and then masked. Segregation performance was evaluated along the horizontal meridian up to retinal eccentricities of about 10 deg. Data are reported from two experiments with different line densities (Kehrer 1989) and two experiments with different orientation contrasts between target lines and background lines (Kehrer 1990). Segregation performance proved to depend strongly on these texture variations, and it peaked several degrees from fixation in all cases. The filter model provided satisfactory predictions of experimental data when model parameters were adjusted appropriately. It is concluded (1) that filter models defined in strictly spatial terms (i.e., without temporal properties) offer a sufficient framework to account for the psychophysical data and (2) that the particular course of the performance curve (i.e., the performance peak outside the central region) must be attributed to the characteristics of second-layer filters. Received: 28 June 2001 / Accepted in revised form: 10 October 2002 / Published online: 13 February 2003 Correspondence to: L. Kehrer (e-mail: lothar.kehrer@uni-bielefeld.de) Acknowledgements. This work was supported by Grant Ke 388/3-2 from the Deutsche Forschungsgemeinschaft (DFG). We wish to thank Jonathan Harrow for improving the English text and an anonymous reviewer for many constructive comments.     

Delayed feedback and multiple attractors in a host–parasitoid system

 Continuous-time, age structured, host–parasitoid models exhibit three types of cyclic dynamics: Lotka–Volterra-like consumer-resource cycles, discrete generation cycles, and “delayed feedback cycles” that occur if the gain to the parasitoid population (defined by the number of new female parasitoid offspring produced per host attacked) increases with the age of the host attacked. The delayed feedback comes about in the following way: an increase in the instantaneous density of searching female parasitoids increases the mortality rate on younger hosts, which reduces the density of future older and more productive hosts, and hence reduces the future per head recruitment rate of searching female parasitoids. Delayed feedback cycles have previously been found in studies that assume a step-function for the gain function. Here, we formulate a general host–parasitoid model with an arbitrary gain function, and show that stable, delayed feedback cycles are a general phenomenon, occurring with a wide range of gain functions, and strongest when the gain is an accelerating function of host age. We show by examples that locally stable, delayed feedback cycles commonly occur with parameter values that also yield a single, locally stable equilibrium, and hence their occurrence depends on initial conditions. A simplified model reveals that the mechanism responsible for the delayed feedback cycles in our host–parasitoid models is similar to that producing cycles and initial-condition-dependent dynamics in a single species model with age-dependent cannibalism. Received: 24 October 1997 / Revised version: 13 June 1998     

An anatomical study of anther development in Acorus L.: Phylogenetic implications

 Critical morphological synapomorphies have not been found in support of the Acoranan hypothesis, the molecular phylogenetic discovery that Acoranae are the basal monocots. The previously undetermined pattern of anther wall development in Acorus has been suggested to be one such character. Two main types of anther wall development have been recognized: 1) the “monocotyledonous” type, which characterizes both monocots and dicots, and 2) the “dicotyledonous” type, which is almost exclusively found among dicots. An anatomical study of anther wall development in Acorus was here undertaken using the electron microscope. Development of the anther wall in Acorus was found to be somewhat irregular or perhaps even intermediate between the two types although largely consistent with the “monocotyledonous” type. The presumed significance of anther wall development and other critical morphological characters to the Acoranan hypothesis in the absence of knowledge about the sister group to the monocots is evaluated. Received August 28, 2000 Accepted February 19, 2001     

Salt tolerance in Lycopersicon spp. VII. Pleiotropic action of genes controlling earliness on fruit yield

 The change from vegetative to reproductive development (earliness) in Lycopersicon chesmannii line L2 was delayed for 20 weeks when compared to other Lycopersicon species under greenhouse conditions. The interspecific hybrid of L. chesmannii L2 and L. esculentum E9, a cherry tomato cultivar, also showed this delay in reproductive development. The distribution of this character in the F₂-derived population showed a bimodal shape, plants could be scored easily as “early” or “late” in two nutrient conditions (optimum and high salinity). A QTL with major effects on earliness was detected in salinity, which explained 35.6% of the phenotypic variation. The effect of this QTL greatly diminished under control conditions, indicating differences in the genetic control of earliness between treatments. ACC synthase or phytochrome B2 are the products of candidate genes for such a major QTL. Other QTLs with minor effects, and epistatic interactions, are also involved in earliness under both conditions. A “late” F₂ subpopulation yielded twice as much as an “early” F₂; conversely, “early” plants were taller than “late” plants, regardless of the treatment. QTL analysis, carried out in both subpopulations, showed that yield differences may be explained by chesmannii alleles showing negative additive effects at some QTLs only in the “early” subpopulation. The effect of population subdivision on QTL analysis was investigated by computer simulations to show sample-size or random effects; thus, important pleiotropic or regulatory effects of genes controlling earliness on yield that affect QTL analysis, have been reveiled. Therefore alleles controlling earliness in L. chesmannii have to be taken into account for a more efficient utilization of the genetic resources of this species. Received: 30 June 1998 / Accepted: 31 August 1998     

Question 1: Commentary Referring to the Statement “The Origin of Life can be Traced Back to the Origin of Kinetic Control” and the Question “Do You Agree with this Statement; and How Would You Envisage the Prebiotic Evolutionary Bridge Between Thermodynamic and Kinetic Control?” Stated in Section 1.1.

The paper summarizes thoughts induced by some of the programmatic questions Pier Luigi Luisi posed for discussion at the Erice workshop. Presented at: International School of Complexity – 4th Course: Basic Questions on the Origins of Life; “Ettore Majorana” Foundation and Centre for Scientific Culture, Erice, Italy, 1–6 October 2006.     

Modeling species richness controlled by community-intrinsic and community-extrinsic processes: coastal fish communities as an example

 This article focuses on the analysis of coastal fish communities along the Norwegian Skagerrak coast. Species numbers are estimated based on annual samples of the fish communities within 12 fjords from 1953 to 1994. On this basis, a community dynamics model (incorporating both community-intrinsic and community-extrinsic processes) was developed and analyzed. This model is then discussed on the basis of other community models available through the literature, both phenomenologically oriented and process-oriented models. Received: January 17, 2002 / Accepted: May 13, 2002 Acknowledgments We thank Dr. Masakado Kawata for the invi-tation to present this paper at the 19th Symposium of the Society of Population Ecology held in Yamagata, Japan, October 26–28, 2001: “Evolution of Biodiversity: Theories and Facts.” Valuable input was provided after the presentation at this meeting, which we greatly appreciated. The reformulation of the model in terms of ΔS was kindly suggested to us by Prof. Joan Roughgarden. Thanks to Dr. Hildegunn Viljugrein for advice on the BUGS analyses and to two anonymous reviewers for constructive comments. This work has been supported by grants from the Norwegian Science Council (NFR). Correspondence to:N.C. Stenseth