Developmental Conversion Research Articles

LIFE CYCLE-ASSOCIATED CHARACTER VARIATION IN AULACOSEIRA KRAMMERI SP. NOV., A NEW MIOCENE SPECIES FROM OREGON, U.S.A.

A new freshwater species, Aulacoseira krammeri R.K. Edgar, Kociolek & S.M. Edgar, from Miocene deposits in Oregon is described. The variation within this species encompasses distinctive robust and gracile morphotypes, and is described emphasizing variation incontinuous characters of valve morphology over the entire vegetative life cycle, especially with respect to covariation of mantle height, interareolar distance (within striae), interstriae distance, ringleist depth and height, and number of ringleist rimoportular canals with size(i.e. valve diameter). The morphometric differentiation of A. krammeri, A. solida (Eulenstein in Van Heurck) Krammer and A. paucistriata Bradbury is examined with an emphasis on size-standardised comparisons and is shown to be largely quantitative. Differences among species in principal axis regression slopes of characters versus valve diameter or mantle height and the location of bivariate means (character and valve diameter or mantle height) were statistically assessed using confidence intervals and confidence ellipses, respectively. A. krammeri displays a unique, non-overlapping distribution of linking and separation spine morphologies with respect to valve diameter and intermediate spine morphologies in a narrow range of cell sizes closely associated with resting spore formation, reflecting a process of developmental conversion.

Expression of Protein Kinase C Inhibitor Blocks Cerebellar Long-Term Depression without Affecting Purkinje Cell Excitability in Alert Mice

A longstanding but still controversial hypothesis is that long-term depression (LTD) of parallel fiber-Purkinje cell synapses in the cerebellum embodies part of the neuronal information storage required for associative motor learning. Transgenic mice in which LTD is blocked by Purkinje cell-specific inhibition of protein kinase C (PKC) (L7-PKCI mutants) do indeed show impaired adaptation of their vestibulo-ocular reflex, whereas the dynamics of their eye movement performance are unaffected. However, because L7-PKCI mutants have a persistent multiple climbing fiber innervation at least until 35 d of age and because the baseline discharge of the Purkinje cells in the L7-PKCI mutants is unknown, factors other than a blockage of LTD induction itself may underlie their impaired motor learning. We therefore investigated the spontaneous discharge of Purkinje cells in alert adult L7-PKCI mice as well as their multiple climbing fiber innervation beyond the age of 3 months. We found that the simple spike and complex spike-firing properties (such as mean firing rate, interspike interval, and spike count variability), oscillations, and climbing fiber pause in the L7-PKCI mutants were indistinguishable from those in their wild-type littermates. In addition, we found that multiple climbing fiber innervation does not occur in cerebellar slices obtained from 3- to 6-month-old mutants. These data indicate (1) that neither PKC inhibition nor the subsequent blockage of LTD induction disturbs the spontaneous discharge of Purkinje cells in alert mice, (2) that Purkinje cell-specific inhibition of PKC detains rather than prevents the developmental conversion from multiple to mono-innervation of Purkinje cells by climbing fibers, and (3) that as a consequence the impaired motor learning as observed in older adult L7-PKCI mutants cannot be attributable either to a disturbance in the baseline simple spike and complex spike activities of their Purkinje cells or to a persistent multiple climbing fiber innervation. We conclude that cerebellar LTD is probably one of the major mechanisms underlying motor learning, but that deficits in LTD induction and motor learning as observed in the L7-PKCI mutants may only be reflected in differences of the Purkinje cell signals during and/or directly after training.

Long-term depression of the cerebellar climbing fiber--Purkinje neuron synapse.

In classic Marr-Albus-Ito models of cerebellar function, coactivation of the climbing fiber (CF) synapse, which provides massive, invariant excitation of Purkinje neurons (coding the unconditioned stimulus), together with a graded parallel fiber synaptic array (coding the conditioned stimulus) leads to long-term depression (LTD) of parallel fiber-Purkinje neuron synapses, underlying production of a conditioned response. Here, we show that the supposedly invariant CF synapse can also express LTD. Brief 5 Hz stimulation of the CF resulted in a sustained depression of CF EPSCs that did not spread to neighboring parallel fiber synapses. Like parallel fiber LTD, CF LTD required postsynaptic Ca2+ elevation, activation of group 1 mGluRs, and activation of PKC. CF LTD is potentially relevant for models of cerebellar motor control and learning and the developmental conversion from multiple to single CF innervation of Purkinje neurons.

Some problems of crustacean taxonomy related to the phenomenon of Horohalinicum

If gene regulatory processes form the basis of phenotypic plasticity that evidences itself as discrete adaptive norms (sensu developmental conversion), it seems evident that the forms capable of such adaptations would possess a correspondingly richer genome than those lacking such adaptations. We examine evidence for the existence of genetic triads which are comprised of one form with wide niches due to phenotypic plasticity and two forms, each which is genetically fixed and exhibits a phenotype at one of the extremes of the plastic form. The invasion of organisms into new environments may occur by acquisition of discrete adaptive norms (DAN), which enable a transition between the old and new environments. The horohalinicum (=critical salinity) is defined as a salinity zone which always corresponds to 5–8 ppm and divides freshwater and marine faunas and floras, as well as many other physico-chemical characteristics. The current paper explores the existence of discrete adaptive norms and genetic triads with respect to salinity in some euryhaline crustaceans, particularly the arctic copepods, and discusses how such adaptations might inform us about species boundaries that affect the taxonomic status of these forms.

Developmental conversion of inotropism by endothelin I and angiotensin II from positive to negative in mice

Inotropic effects on isolated neonatal and adult mouse myocardium of endothelin I and angiotensin II were examined. Endothelin I produced a sustained positive inotropic response in the neonate but a sustained negative response in the adult. Both were concentration-dependent and were inhibited by the endothelin ET A receptor antagonist, BQ-123 (Cyclo( d-a-aspartyl- l-prolyl- d-valyl- l-leucyl- d-tryptophyl)). Angiotensin II produced a sustained positive inotropic response in the neonate while a sustained negative response in the adult. Both were concentration-dependent and were inhibited by the angiotensin AT 1 receptor antagonist, YM358 (2,7-diethyl-5-((2′-(1 H-tetrazol-5-yl)biphenyl-4-yl)methyl-5 H-pyrazolo(1,5-b)(1,2,4)triazole potassium salt monohydrate). These results indicate that inotropic responses of the mouse heart to cardioactive peptides are unique among experimental animal species and may be reversed during development.

Towards a more synthetic view of evolution

The 21st century will be the century of biology, and evolutionary biology will be the linchpin. Yes, this is a bold statement, but it is certainly in keeping with Dobzhansky's famous dictum about evolution. Dobzhansky was one of the major architects of the Modern Synthesis. This book by Schlichting and Pigliucci (SP the Final Synthesis is just gathering steam. Note that these dates are just rough guides and not meant to imply that significant work did not occur outside those periods. What is critical is that within these periods the relevant disciplines all become focussed on the same research program. We are now in the midst of this final joining of biological disciplines. This joining, if history is a guide, will take the next couple of decades. This joining will be driven from the evolutionary side. It is the evolutionary side within which the disciplines are linked by a synthetic theory. The molecular side is primarily linked by a shared set of techniques and a viewpoint that centers all questions ultimately on DNA sequences. Because the evolution side is much more theory driven, that is where synthetic concepts tend to arise. This book by S&P is an excellent example of such theorydriven synthesis. Also, if truth be told, as a whole evolutionary biologists tend to have a larger perspective than those in many other biological disciplines. Looking back on the Modern Synthesis is useful for the task ahead (Mayr and Provine, 1980). In particular, if we learn anything from that episode in our history, we

The Developmental Conversation: Facilitating Moral and Intellectual Growth in Our Students

AbstractWhereas the legalistic framework is defined by institutional rules and standards, policies and procedures focusing on students as learners and as agents of their own development should be considered as a new framework for student affairs practitioners.

Sweet pepper plastids: enzymic equipment, characterisation of the plastidic oxidative pentose-phosphate pathway, and transport of phosphorylated intermediates across the envelope membrane

Chloroplasts or chromoplasts were purified from sweet-pepper (Capsicum annuum L. cv. Yolo Wonder) fruits and analysed with respect to their enzymic equipment, the transport properties across the envelope membrane, and for the presence of a functional oxidative pentose-phosphate pathway (OPPP). It was demonstrated that both types of plastid contain enzyme activities that allow glycolysis and OPPP. During the developmental conversion from chloroplasts to chromoplasts the activities of enzymes catalysing potentially rate-limiting reactions in glycolysis increased considerably. Most enzyme activities involved in the plastidic OPPP stayed constant or decreased during ripening, but transaldolase activity increased by more than 500%. To analyse whether pepper fruit chromoplasts are able to use exogenously supplied carbohydrates for the OPPP we measured the rate of 14CO2 release after application of radioactively labelled precursors. Isolated pepper fruit chromoplasts used exogenously supplied [U14C]glucose- 6-phosphate (Glc6P) as a precursor for the OPPP. The metabolic flux through this pathway was stimulated by the presence of additional compounds which require reducing equivalents for further conversion, e.g. nitrite, or 2-oxoglutarate plus glutamine. The [14C]Glc6P-driven OPPP in isolated chromoplasts exhibited saturation with rising concentrations of Glc6P, reaching highest rates at an external concentration of about 2 mM. Exogenously given [U14C]glucose 1-phosphate (Glc1P)′ did not lead to a release of 14CO2, indicating that this hexose phosphate is not taken up into the intact plastid. Using a proteoliposome system in which the envelope membrane proteins from sweet-pepper chromoplasts were functionally reconstituted we demonstrated that Glc6P is transported in counter-exchange with inorganic phosphate (Pi) or other phosphorylated intermediates. The Glc6P was taken up into proteoliposomes with an apparent Km of 0.34 mM. Surprisingly, in contrast to tomato fruit plastids, isolated chromoplasts from sweet-pepper fruits do not possess a phosphate translocator allowing the uptake of Glc1P. Rising exogenous concentrations of dihydroxyacetone phosphate strongly inhibited the metabolic flux through the OPPP. This observation is discussed with respect to the presence of two phosphate translocator proteins in the envelope of sweet-pepper chromoplasts and with respect to possible metabolic changes occurring in heterotrophic tissues during development.

Effect of sympathectomy on inotropic responsiveness to alpha-adrenoceptor stimulation in developing mouse myocardia.

Effects of postnatal sympathectomy on inotropic responsiveness to alpha-adrenoceptor stimulation were examined in mouse myocardia to determine whether the developmental conversion of alpha-adrenoceptor-mediated inotropic responses from positive to negative is triggered by sympathetic innervation. Sympathectomy was performed chemically by consecutively administering 6-hydroxydopamine for 14 days after birth and confirmed by the absence of inotropic responses to tyramine. In newborn myocardia, phenylephrine, in the presence of propranolol, produced concentration-dependent positive inotropic responses. Three weeks after birth, phenylephrine, in the presence of propranolol, produced concentration-dependent negative inotropic responses, both in control and in sympathectomized myocardia; no difference was observed between the two groups of mice in the maximum decrease in contractile force produced by phenylephrine. The sensitivity (pD2 value) to phenylephrine was significantly higher in sympathectomized myocardia. In conclusion, sympathetic innervation of the mouse ventricular myocardium is not required for the developmental conversion of the alpha-adrenoceptor-mediated inotropic response from positive to negative.

Gene regulation, quantitative genetics and the evolution of reaction norms

The ideas of phenotypic plasticity and of reaction norm are gaining prominence as important components of theories of phenotypic evolution. Our understanding of the role of phenotypic plasticity as an adaptation of organisms to variable environments will depend on (1) the form(s) of genetic and developmental control exerted on the shape of the reaction norm and (2) the nature of the constraints on the possible evolutionary trajectories in multiple environments. In this paper we identify two categories of genetic control of plasticity: allelic sensitivity and gene regulation. These correspond generally to two classes of response by the developmental system to environmental change: phenotypic modulation, in which plastic responses are a continuous and proportional function of environmental stimuli and developmental conversion, where responses tend to be not simply proportional to the stimuli. We propose that control of plasticity by regulatory actions has distinct advantages over simple allelic sensitivity: stability of phenotypic expression, capacity for anticipatory response and relaxation of constraints due to genetic correlations. We cite examples of the extensive molecular evidence for the existence of environmentally-cued gene regulation leading to developmental conversion. The results of quantitative genetic investigations on the genetics and evolution of plasticity, as well as the limits of current approaches are discussed. We suggest that evolution of reaction norms would be affected by the ecological context (i.e. spatial versus temporal variation, hard versus soft selection, and fine versus coarse environmental grain). We conclude by discussing some empirical approaches to address fundamental questions about plasticity evolution.

Primary, self-renewing erythroid progenitors develop through activation of both tyrosine kinase and steroid hormone receptors.

Self renewal in the hematopoietic system is thought to be restricted to a class of pluripotent stem cells. The capacity of cells with the properties of committed progenitors to self renew in many leukemias is thought to be an abnormal property resulting from the mutations responsible for leukemic transformation. It is not known how cells that can self-renew differ from cells that cannot. The notion that only pluripotent stem cells self renew has recently been challenged: normal committed erythroid progenitors capable of sustained self renewal have been described. These cells, called SCF/TGF alpha progenitors, co-express the c-Kit receptor tyrosine kinase and c-ErbB, the avian receptor for epidermal growth factor and transforming growth factor (TGF) alpha, and they undergo continuous self renewal in response to TGF alpha and estradiol. In contrast, common erythroid progenitors (termed SCF progenitors) express only c-Kit and undergo a limited number of cell divisions in response to the c-Kit ligand, stem cell factor (SCF). Both types of progenitor faithfully reproduce terminal erythroid differentiation in vitro when exposed to differentiation factors. Here, we have investigated the developmental origin of these two classes of self-renewing erythroid progenitors. We show that SCF progenitors can develop into SCF/TGF alpha progenitors. This developmental conversion requires 10-14 days and is accompanied by a gradual up-regulation of bioactive TGF alpha receptor. Using sera depleted of endogenous growth factors, we demonstrate that the development of SCF progenitors into SCF/TGF alpha progenitors absolutely requires the simultaneous presence of SCF, TGF alpha and estradiol, and is strongly enhanced by an unknown activity in chicken serum. SCF progenitors can be induced to develop into self-renewing SCF/TGF alpha progenitors. The development of self renewal is triggered by specific combinations of growth factors and hormones. This has important implications for understanding leukemogenesis, as the self renewal of leukemic cells may reflect the normal potential of certain committed progenitor cells and not, as has been thought, a unique abnormal property of leukemic cells.

Sustained negative inotropism mediated by α‐adrenoceptors in adult mouse myocardial developmental conversion from positive response in the neonate

1. Inotropic responses to alpha-adrenoceptor stimulation and the effects of antagonists were examined in isolated ventricular preparations from neonatal and adult mice. 2. Phenylephrine, in the presence of propranolol, produced positive inotropic responses in neonates up to 1 week after birth, while it produced negative inotropic responses in mice older than 3 weeks. 3. Both positive and negative responses to phenylephrine in neonates and adults, respectively, were antagonized by prazosin, WB4101 (2-([2,6-dimethoxyphenoxyethyl]aminomethyl)-1,4-benzodioxane) and 5-methylurapidil, but not by atropine, yohimbine or chlorethylclonidine. 4. Noradrenaline (NA) produced positive inotropic responses both in the neonate and adult; the responses were observed in a lower concentration-range in the neonate than in the adult. WB4101 produced a significant leftward shift of the concentration-response curve for noradrenaline in adult preparations while only a slight rightward shift was observed in the neonate. 5. Our results demonstrate the presence of alpha-adrenoceptor-mediated inotropic responses in the mouse ventricular myocardia. The response to phenylephrine changes from a positive to a negative effect during postnatal development. The responses are mediated by alpha 1-adrenoceptors, and modulate the overall inotropic response to NA in the adult.

P1-178 - Developmental conversion of alpha-adrenoceptor-mediated inotropism from positive to negative in mouse myocardium is not inhibited by postnatal sympathetic denervation

P1-178 - Developmental conversion of alpha-adrenoceptor-mediated inotropism from positive to negative in mouse myocardium is not inhibited by postnatal sympathetic denervation

P-314 - Alpha adrenoceptor-mediated inotropism in mouse myocardia: developmental conversion from positive to negative effects.

P-314 - Alpha adrenoceptor-mediated inotropism in mouse myocardia: developmental conversion from positive to negative effects.

Identification of a novel large extrachromosomal DNA (LED) in the Trypanosomatidae.

We have identified a novel 75 kbp large extrachromosomal DNA (LED) which is stably maintained during developmental conversion of Trypanosoma cruzi. It has a covalently closed circular conformation and is not derived from the kinetoplast network. In all T. cruzi strains analysed, LED contains 18S rRNA and spliced leader (sl) sequences. LED from the T. cruzi Y strain contains a minimum of 15 copies of the sl repeat arrayed in a head-to-tail configuration and 50 copies of a 196 bp repeat. LED is also present in Trypanosoma dionisii (subgenus Schizotrypanosoma) and in other members of the family Trypanosomatidae. LED from different T. cruzi strains and from other members of the Trypanosomatidae differ in their content of large ribosomal subunit rRNA sequences and the 196 bp repeat. The presence of LED in four evolutionarily distant trypanosomatid species suggests that it plays an important role in the biology of these parasites.

Role of calcium and protein kinase C in development of the delayed rectifier potassium current in xenopus spinal neurons

The delayed rectifier current of embryonic Xenopus spinal neurons plays the central role in developmental conversion of calcium-dependent action potentials to sodium-dependent spikes. During its maturation, this potassium current undergoes a pronounced increase in rate of activation. The mechanism underlying the change in kinetics was analyzed with whole-cell voltage clamp of neurons cultured under various conditions. Calcium is necessary at an early stage of development, to permit influx that triggers subsequent release of calcium from intracellular stores. Its action is prevented by depletion of protein kinase C and mimicked by stimulation of the kinase. Calcium influx through voltage-dependent channels at early stages of development regulates the differentiation of potassium current kinetics and modulation of the ionic dependence of action potentials.

Arrest of developmental conversion of type II to type I fibres after suspension hypokinesia

The effects of hypokinesia and of the lack of gravity on muscle fibres, fibre type composition and myosin light chain pattern, as well as on muscle mechanoreceptors were investigated in the slow-twitch soleus (SOL) and fast-twitch extensor digitorum longus (EDL) muscles of young growing and adult rats after suspension hypokinesia (SH) of their hind limbs. The animals were suspended by their tail so that their hind limbs were relieved of their normal weight-bearing function for 3-6 weeks. In normal 3- to 4-week-old rats the SOL contained about 50% type I fibres and their percentage increased up to about 80% until the 10th week, with simultaneous reduction of type IIA fibres. After 3 to 6 weeks of suspension treatment maintained from 3- to 4-week-old rats up to 6 to 10 weeks of age, the SOL still only contained about 50% of type I fibres. The content of fast LC1 and LC2 in the SOL of 6-week-old rats after 3 weeks of suspension was higher than that of control litter-mates reflecting the higher occurrence of IIA fibres in the suspended solei. No changes in fibre type composition were observed after SH performed in adult rats. SH thus leads, in young animals, to the arrest of conversion of type IIA to type I fibres resulting in the persistence of the fibre type composition and of the myosin light chain pattern corresponding to those present in the SOL at the time of the onset of suspension. In both young and adult rats, SH markedly decreased the mass and the mean cross-sectional area of the SOL, mainly due to the severe atrophy of type I fibres. We observed no signs indicating conversion of type I back to type IIA muscle fibres due to the SH either in young or adult animals. In contrast to profound changes in the SOL, no significant differences were found in the EDL in any of the parameters studied. No changes in the investigated parameters of muscle spindles and tendon organs were observed after SH, performed either in young or in adult rats. We thus conclude that SH leads to muscle atrophy and that it influences mainly or exclusively type I fibres in muscles with a postural function such as the SOL.(ABSTRACT TRUNCATED AT 400 WORDS)

Genomic variation of Trypanosoma cruzi: involvement of multicopy genes

By using improved pulsed field gel conditions, the karyotypes of several strains of the protozoan parasite Trypanosoma cruzi were analyzed and compared with those of Leishmania major and two other members of the genus Trypanosoma. There was no difference in chromosome migration patterns between different life cycle stages of the T. cruzi strains analyzed. However, the sizes and numbers of chromosomal bands varied considerably among T. cruzi strains. This karyotype variation among T. cruzi strains was analyzed further at the chromosomal level by using multicopy genes as probes in Southern hybridizations. The chromosomal location of the genes encoding alpha- and beta-tubulin, ubiquitin, rRNA, spliced leader RNA, and an 85-kilodalton protein remained stable during developmental conversion of the parasite. The sizes and numbers of chromosomes containing these sequences varied among the different strains analyzed, implying multiple rearrangements of these genes during evolution of the parasites. During continuous in vitro cultivation of T. cruzi Y, the chromosomal location of the spliced leader gene shifted spontaneously. The spliced leader gene encodes a 35-nucleotide RNA that is spliced in trans from a 105-nucleotide donor RNA onto all mRNAs in T. cruzi. The spliced leader sequences changed in their physical location in both the cloned and uncloned Y strains. Associated with the complex changes was an increase in the infectivity of the rearranged variant for tissue culture cells. Our results indicate that the spliced leader gene clusters in T. cruzi undergo high-frequency genomic rearrangements.

Canalization Versus Developmental Conversion in a Spatially Variable Environment

A model is presented for the evolution of developmental control in a spatially variable environment. Individuals are assumed to disperse at random into one of two patches (one harsh and the other benign) and use one of three developmental strategies for the production of one of two discrete morphological types. Two of the strategies are unconditional (develop as either the stress-tolerant or the nontolerant morph), and the third strategy depends on the environment. Invasion criteria are used to determine the conditions under which each of the three pure strategies are evolutionarily stable and the conditions under which the population is expected to contain some mixture of these strategies at equilibrium. The results demonstrate that environmental control of development requires a cost to the stress-tolerant morphology, and that the average probability of making the right choice is greater than 50%. The range of patch frequencies over which environmental control is stable increases with increases in these...

Developmental Plasticity: Developmental ConversionversusPhenotypic Modulation

The developmental mechanisms by which the environment may alter the phenotype during development are reviewed. Developmental plasticity may be of two forms: developmental conversion or phenotypic modulation. In developmental conversion, organisms use specific environmental cues to activate alternative genetic programs controlling development. These alternative programs may either lead to alternative morphs, or may lead to the decision to activate a developmental arrest. In phenotypic modulation, nonspecific phenotypic variation results from environmental influences on rates or degrees of expression of the developmental program, but the genetic programs controlling development are not altered. Modulation, which is not necessarily adaptive, is probably the common form of environmentally induced phenotypic variation in higher organisms, and adaptiveness of phenotypic plasticity therefore cannot be assumed unless specific genetic mechanisms can be demonstrated. The genetic mechanisms by which developmental plasticity may evolve are reviewed, and the relationship between developmental plasticity and evolutionary plasticity are examined.