Small Tadpoles Research Articles

Stabilizing massless fields with fluxes in Landau-Ginzburg models

Recent work on flux compactifications suggests that the tadpole constraint generically allows only a limited number of complex structure moduli to become massive, i.e., be stabilized at quadratic order in the spacetime superpotential. We study the effects of higher-order terms systematically around the Fermat point in the 19 Landau-Ginzburg model. This model lives at strong coupling and features no Kähler moduli. We show that indeed massless fields can be stabilized in this fashion. We observe that, depending on the flux, this mechanism is more effective when the number of initially massless fields is large. These findings are compatible with both the tadpole conjecture and the massless Minkowski conjecture. Along the way, we complete the classification of integral flux vectors with small tadpole contribution. Thereby we are closing in on a future complete understanding of all possible flux configurations in the 19 Landau-Ginzburg model.

Effects of metolachlor and fish cues on tadpole antipredator behaviors

Antipredator behaviors are often crucial for animal survival. In environments impacted by humans, animals may be exposed to a wide array of chemical contaminants due to agriculture, industry, and other activities. Among other effects, these chemical contaminants may influence behaviors prey animals use to cope with predation risk, likely reducing survival by suppressing or even maladaptively reversing normal behavioral responses. Changes in behavioral responses could be the result of impaired detection of cues that indicate predator presence or altered behavioral plasticity in response to predator cues. In a laboratory experiment, we tested the effects of the commonly used herbicide metolachlor on the behavior of tadpoles of the American bullfrog (Lithobates catesbeianus) in both the absence and presence of water conditioned with cues (odor) of fish. We observed several behaviors in the tadpoles reflective of predator avoidance when exposed to combinations of metolachlor (0 or 80 ppb) and the presence or absence of fish cues. We found that metolachlor had size-dependent effects on tadpole activity levels, with larger tadpoles being much less active in the absence of metolachlor but similar in activity to small tadpoles when metolachlor was present. Fish cues similarly had size-dependent effects on multiple tadpole behaviors. Finally, hiding behavior and zone change (movement) behavior of tadpoles in response to fish cues varied depending on the presence of metolachlor and, in the case of zone changes, tadpole size. Our findings indicate that the presence of chemical contaminants has the potential to alter important antipredator behaviors. However, these effects are complex, and size may mediate the nature of the response to both chemical contaminants and predators.

The tadpole conjecture in the interior of moduli space

We revisit moduli stabilization on Calabi-Yau manifolds with a discrete symmetry. Invariant fluxes allow for a truncation to a symmetric locus in complex structure moduli space and hence drastically reduce the moduli stabilization problem in its dimensionality. This makes them an ideal testing ground for the tadpole conjecture. For a large class of fourfolds, we show that an invariant flux with non-zero on-shell superpotential on the symmetric locus necessarily stabilizes at least 60% of the complex structure moduli. In case this invariant flux induces a relatively small tadpole, it is thus possible to bypass the bound predicted by the tadpole conjecture at these special loci. As an example, we discuss a Calabi-Yau hypersurface with h3,1 = 3878 and show that we can stabilize at least 4932 real moduli with a flux that induces M2-charge Nflux = 3.

Symmetric fluxes and small tadpoles

The analysis of type IIB flux vacua on warped Calabi-Yau orientifolds becomes considerably involved for a large number of complex structure fields. We however show that, for a quadratic flux superpotential, one can devise simplifying schemes which effectively reduce the large number of equations down to a few. This can be achieved by imposing the vanishing of certain flux quanta in the large complex structure regime, and then choosing the remaining quanta to respect the symmetries of the underlying prepotential. One can then implement an algorithm to find large families of flux vacua with a fixed flux tadpole, independently of the number of fields. We illustrate this approach in a Calabi-Yau manifold with 51 complex structure moduli, where several reduction schemes can be implemented in order to explicitly solve the vacuum equations for that sector. Our findings display a flux-tadpole-to-stabilized-moduli ratio that is marginally above the bound proposed by the Tadpole Conjecture, and we discuss several effects that would take us below such a bound.

Relevant dilaton stabilization

We propose a simple modification of the Goldberger-Wise mechanism for stabilizing the scale of spontaneously broken conformal theories. The source of explicit conformal symmetry breaking is a relevant operator with a small coefficient, as opposed to the usual mechanism of an almost marginal operator with an order-one coefficient. In the warped 5D picture this relevant stabilization corresponds to a small tadpole for the bulk scalar on the UV brane, which can be technically natural if it is the only source for the breaking of a symmetry (for example, a discrete Z2). This modification of the stabilization mechanism has significant consequences for the nature of the conformal phase transition, since the radion/dilaton potential is no longer shallow. The bounce action is significantly reduced, leading to a weaker first-order phase transition instead of the supercooled and strongly first-order transition seen in Goldberger-Wise stabilization. This also leads to reduction of gravitational wave signals which, however, may still be observable at future detectors. We present numerical and analytical studies of the phase transition and the resulting gravitational wave signal strength, assuming that the effective dilaton potential provides a good leading approximation. While the dilaton is not expected to be generically light in this setup, in order to keep perturbative control over the effective theory one needs to mildly tune the dilaton quartic to be somewhat small.

Plasticity for the kin and conspecific preferences in the frog tadpoles (Rana ornativentris).

In vertebrates, little is known of kin recognition systems and their plasticity. Even in well-studied anuran larvae (tadpoles), the determinants and effects of prior experience have not been clarified. This study evaluates the plasticity of kin and conspecific discrimination in tadpoles of the Japanese montane brown frog Rana ornativentris. We raised tadpoles under two different sibship conditions: the pure line, comprising only siblings, and the mixed line, comprising both siblings and non-siblings. The association preference by a subject tadpole to unfamiliar ("stimulus") tadpoles was assessed through binary-choice tests using a 2 × 2 × 2 factorial design among each kinship line (pure and mix), subject ontogeny/size (early stage/small and late-stage/large), and stimuli ontogeny/size. Contrary to our expectations, kin preference was confirmed only in early developmental small tadpoles from mixed line, and only with a small stimulus. Furthermore, tadpoles from mixed line did not exhibit size preference for unrelated conspecifics. These results suggest that different prior associations have modulated kin templates along tadpole ontogeny and that the presence of non-kin would enhance the learning of kin/non-kin. This study provides the first example that plasticity of kin recognition affects not only kin-biased association but also conspecific recognition along ontogeny in tadpoles.

The mechanics of air breathing in African clawed frog tadpoles, Xenopus laevis (Anura: Pipidae).

Frog larvae (tadpoles) undergo many physiological, morphological and behavioral transformations throughout development before metamorphosing into their adult form. The surface tension of water prevents small tadpoles from breaching the surface to breathe air (including those of Xenopus laevis), forcing them to acquire air using a form of breathing called bubble sucking. With growth, tadpoles typically make a behavioral/biomechanical transition from bubble sucking to breaching. Xenopus laevis tadpoles have also been shown to transition physiologically from conforming passively to ambient oxygen levels to actively regulating their blood oxygen. However, it is unknown whether these mechanical and physiological breathing transitions are temporally or functionally linked, or how both transitions relate to lung maturation and gas exchange competency. If these transitions are linked, it could mean that one biomechanical breathing mode (breaching) is more physiologically proficient at acquiring gaseous oxygen than the other. Here, we describe the mechanics and development of air breathing and the ontogeny of lung morphology in X. laevis throughout the larval stage and examine our findings considering previous physiological work. We found that the transitions from bubble sucking to breaching and from oxygen conforming to oxygen regulation co-occur in X. laevis tadpoles at the same larval stage (Nieuwkoop-Faber stages 53-56 and 54-57, respectively), but that the lungs do not increase significantly in vascularization until metamorphosis, suggesting that lung maturation, alone, is not sufficient to account for increased pulmonary capacity earlier in development. Although breach breathing may confer a respiratory advantage, we remain unaware of a mechanistic explanation to account for this possibility. At present, the transition from bubble sucking to breaching appears simply to be a consequence of growth. Finally, we consider our results in the context of comparative air-breathing mechanics across vertebrates.

Re-description of external morphology and factors affecting body and tail shape of the stone frog tadpoles

Ecological studies testing the preponderance of environmental filters on ontogeny to explain the variation in tadpole morphology are scarce for Neotropical anurans. We used tadpoles of the stone frog Limnomedusa macroglossa (Alsodidae): (1) to assess the variation in body and tail shape; (2) to examine the effect of streamlet depth and allometry on tadpole shape, and (3) to re-describe and compare the tadpole external morphology with closely related species. We obtained the body shape and size from 150 tadpoles. The re-description was based on 57 qualitative and 24 quantitative characters, from 19 tadpoles between stages 30 and 37 and 31 to 37, respectively. Allometry was the major factor influencing the lateral view of body shape: smaller tadpoles had round bodies and eyes and nostrils positioned more laterally in comparison with larger ones. Thus, the power of ontogenetic variations reported here makes the tadpole developmental “climax” period a questionable concept that deserves additional attention. The depth gradient of streamlets also affected the shape: in shallower environments, the tadpoles presented a decrease in height of the body, fins and tail muscles, and an increase in body width. These results may indicate adaptations allowing better swimming performance in lotic environments with intense water flow. The external morphological characterization of L. macroglossa presented here differed from that previously reported, mainly due to coloration, body shape, nostril, anal tube, tail, shape and position of nostrils and snout. Additionally, we presented unknown traits for this species, making comparisons with closely related species within the Alsodidae family.

Thermal and salinity effects on locomotor performance of Thoropa taophora tadpoles (Anura, Cycloramphidae).

It is well known that environmental temperature influences several biological functions of ectotherms, notably in amphibians. The high permeability of anuran skin, associated with the effect of elevated environmental temperature, potentiates the dehydration process and this combination may restrict locomotor performance. Thoropa taophora is an endemic species from the Atlantic Rainforest whose tadpoles are semiterrestrial and predominantly diurnal, and are found in rocky seashores where they are exposed to sea spray and high temperatures. In this study we investigated how temperature and salinity conditions affect the locomotor performance in Thoropa taophora tadpoles. We also assessed how different osmotic concentrations affect the activity of the metabolic pathways that support muscle function. We measured the sprint speed of tadpoles of various sizes at different temperatures and salinities in the field. We also measured the activity of the enzymes pyruvate kinase (PK), lactate dehydrogenase (LDH) and citrate synthase (CS) in different temperatures and osmotic concentrations, and calculated the thermal sensitivity and the activity constants for each osmolality. Our results showed that, in general, sprint speed decreased with increasing temperature and salinity. However, whereas the effect of increased salinity was similar in smaller and larger tadpoles, increased temperature had a higher negative impact on sprint speed of larger tadpoles, thus indicating low thermal sensitivity of small tadpoles. PK and LDH thermal sensitivities and LDH constant of activity decreased as the osmolality increased. In conclusion, the locomotor capacity of tadpoles was decreased by temperature and salinity, which may be related to a decrease in anaerobic metabolism both in terms of sensitivity and total energy turnover through enzymatic activity. We discuss the ecological consequences, including the potential impacts on predator escape behavior promoted by changes in metabolism and locomotor performance in an early stage of development of this species.

Histology and microscopic anatomy of the thyroid gland during the larval development of Pseudis platensis (Anura, Hylidae).

Several hormones regulate anuran larval development, most notably thyroid hormones (THs). In anurans, metamorphosis fails when the thyroid gland is absent or inactivated, resulting in giant tadpoles. Larval gigantism occurs naturally in neotropical frogs of the genus Pseudis as a result of a prolonged larval period. Its thyroid function is poorly investigated and the focus of this study. We describe qualitative and quantitative variations in larval development for field-captured specimens of Pseudis platensis and compare those to the development of two sympatric species, Phyllomedusa sauvagii and Pithecopus azureus, which have small tadpoles and a shorter larval period. We describe morphological changes in the thyroid glands of larval and adult specimens. In contrast to other species with similar ecological requirements, P. platensis exhibits distinct glandular activity. During premetamorphosis, there was little or no thyroid activity, a period in which the tadpole reached 70% of its maximum size. Development and degree of activity of the thyroid gland determine the duration of the early stages of the larval period. Thyroid gland histology in tadpoles appears to correlate with the TH activity, and in turn with the diversity in anuran life history transitions.

Developmental effects on social preferences in frog tadpoles, Rana ornativentris

Because the costs and benefits of living in a group change according to ontogenetic stage, variation in body size is predicted to cause differences in social preferences even within individuals. Based on the fitness consequences of group living, small individuals in early developmental stages are predicted to form aggregations to reduce predation risk, whereas large individuals will avoid one another to reduce resource competition. Furthermore, when individuals recognize kin, social preferences may vary depending on the kinship relationship between them. In this study, we examined the association preference of different-sized Rana ornativentris tadpoles, that is, small individuals in early developmental stages and larger tadpoles at later developmental stages. Our binary choice experiments showed that, regardless of size, R. ornativentris tadpoles can discriminate between conspecifics based on both phenotype and relatedness. That is, association preferences were determined by the size and relatedness of the subject and the two stimulus tadpoles. Both small and large individuals spent longer with small than large tadpoles. However, there was a significant difference in preference between small and large tadpoles. When one of the two stimulus tadpoles was a large nonsibling, small tadpoles tended to associate with both small siblings and small nonsiblings. In contrast, large tadpoles showed no preference for small siblings. In the context of the benefits of grouping, phenotype matching would be effective only in small tadpoles. Conversely, under the pressure of conspecific competition, large tadpoles did not prefer to associate with similar-sized ones; this may have enabled large tadpoles to avoid competing with kin and to reduce the costs of group living.

A new diploid species belonging to the Odontophrynus americanus species group (Anura: Odontophrynidae) from the Espinhaço range, Brazil.

The Odontophrynus americanus species group is a complex of diploid and tetraploid species hardly distinguished by morphological characters. It currently consists of three allopatric diploid species (i.e. O. cordobae, O. lavillai, and O. maisuma) and one widely distributed tetraploid species (i.e. O. americanus). We herein describe a new diploid allopatric species from campo rupestre, a typical phytophysiognomy of the Espinhaço Range, Brazil. The new species is distinguishable by the diploid complement of 2n = 2x = 22 chromosomes, small to medium-sized dorsal dark brown blotches with low contrast on a light brown background, light mid-dorsal stripe absent or greatly interrupted in most specimens with yellowish coloration as the background of both head and flanks of the body, advertisement call with dominant frequency of 840-1080 Hz, pulse rate of 90.5-106.7 pulses/s, and small tadpoles (TL = 24.30-35.69 mm).

Contacts with large, active individuals intensify the predation risk of small conspecifics.

Size variation within a population can influence the structure of ecosystem interactions, because ecological performance differs between individuals of different sizes. Although the impact of size variation in a predator species on the structure of interactions is well understood, our knowledge about how size variation in a prey species might modify the interactions between predators and prey is very limited. Here, by examining the interactions between predatory Hynobius retardatus salamander larvae and their prey, Rana pirica frog tadpoles, we investigated how large prey individuals affect the predation mortality of small prey conspecifics. First, in an experiment conducted in a field pond in which we manipulated the presence of salamanders and large tadpoles (i.e., large enough to protect them against salamander predation) with small tadpoles, we showed that in the presence of large tadpoles the mortality of small tadpoles from salamander predation was increased. On the basis of our observations of the activity of individuals, we hypothesized that active large tadpoles caused physical disturbances, which in turn caused the small tadpoles to move, and thus increased their encounter frequency with the predatory salamanders. To test this hypothesis, we conducted a laboratory experiment in small tanks with three players (i.e., one salamander as predator, one small tadpole as focal prey, and either a small or a large tadpole as the prospective movement inducer). In each tank, we manipulated the presence or absence of a movement inducer, and, when present, its size (large or small) and access (caged or uncaged) to the focal prey. In the presence of a large, uncaged movement inducer, the focal prey was more active and suffered from higher predation mortality compared with the other treatments, because the large movement inducer (unlike a small movement inducer) moved actively and, when uncaged, could stimulate movement of the focal prey through direct contact. The results indicated that high activity of large prey individuals and the resulting behavioral interactions with small conspecifics via direct contact indirectly increased the mortality of the small prey.

Metabolomic approach for identifying and visualizing molecular tissue markers in tadpoles of Xenopus tropicalis by mass spectrometry imaging.

ABSTRACTIn developmental and cell biology it is crucial to evaluate the dynamic profiles of metabolites. An emerging frog model system using Xenopus tropicalis, whose genome sequence and inbred strains are available, is now ready for metabolomics investigation in amphibians. In this study we applied matrix-assisted laser desorption/ionization (MALDI)-mass spectrometry imaging (MSI) analysis to identify and visualize metabolomic molecular markers in tadpoles of Xenopus tropicalis. We detected tissue-specific peaks and visualized their distribution in tissues, and distinguished 19 tissues and their specific peaks. We identified, for the first time, some of their molecular localizations via tandem mass spectrometric analysis: hydrocortisone in artery, L-DOPA in rhombencephalon, taurine in eye, corticosterone in gill, heme in heart, inosine monophosphate and carnosine in muscle, dopamine in nerves, and phosphatidylethanolamine (16:0/20:4) in pharynx. This is the first MALDI-MSI study of X. tropicalis tadpoles, as in small tadpoles it is hard to distinguish and dissect the various organs. Furthermore, until now there has been no data about the metabolomic profile of each organ. Our results suggest that MALDI-MSI is potentially a powerful tool for examining the dynamics of metabolomics in metamorphosis as well as conformational changes due to metabolic changes.

Tadpoles of three common anuran species from Thailand do not prey on mosquito larvae.

Tadpoles are often considered to be predators of mosquito larvae and are therefore beneficial for the control of certain disease vectors. Nevertheless, only a few species have actually been recorded to prey on mosquito larvae. The mosquito larvae predation rates of tadpoles of three common Thai anuran species (Bufo melanostictus, Kaloula pulchra and Hylarana raniceps) were experimentally tested. Tadpoles in varying developmental stages were used to assess a size/age effect on the predation rate. In addition, different instars of Culex quinquefasciatus were used in order to assess a prey size effect on the predation rates. All three species failed to show any evidence of mosquito larvae predation. Neither small nor large tadpoles fed on mosquito larvae. Prey size also did not affect predation. Although tadpoles do not feed on mosquito larvae, there may be other direct or indirect inter-specific interactions that adversely impact the development of larvae in shared habitats with tadpoles.

Antagonistic indirect interactions between large and small conspecific prey via a heterospecific predator

Intrapopulation size variation strongly influences ecological interactions because individuals belonging to different size groups have distinct functions. Most demonstrations of the impacts of size variation in trophic systems have focused on size variation in predator species, and the consequences of size variation in prey species are less well understood. We investigated how prey size structure shapes intra- and interspecific interactions in experiments with a gape-limited predator (larvae of the salamander Hynobius retardatus) and its heterospecific prey (frog tadpoles, Rana pirica). We found that large and small tadpole size groups each increased mortality in the other group by intensifying salamander predation; this type of indirect interactions is called apparent competition. The antagonistic impacts on the prey size groups were caused by different size-specific mechanisms. By consuming small tadpoles, the salamanders grew large enough to consume large tadpoles. The activity of large tadpoles, by increasing the activity of the small tadpoles, may increase the number of encounters with the predator and thus small tadpole mortality. These results suggest that the magnitude of a predator's ecological role, such as whether a top–down trophic cascade is initiated, depends on size variation in its heterospecific prey.

Body size and group size of Cuban tree frog (Osteopilus septentrionalis) tadpoles influence their escape behaviour

Tadpoles risk attack from both aquatic and aerial predators. We investigated how body size and group size influenced the behaviour of tadpoles before and during a predatory attack from above to test the predictions of the theoretical economic escape model. We examined escape (swimming) response of small and large Cuban tree frog (Osteopilus septentrionalis) tadpoles kept under three density treatments and predicted that increased group size, body size and depth in the water column would all reduce perceived risk and, therefore, escape responses to simulated predation. Compared with the lower density groups, tadpoles in higher density groups moved shorter distances, and many individuals did not even move away in response to being touched. Contrary to our predictions based on the economic escape model, smaller tadpoles (which should be more vulnerable to a greater suite of predators) were less reactive than larger tadpoles, and this result may reflect different costs of escape. Finally, although tadpoles might be exposed to a wider range of predator species (aerial as well as aquatic predators), we found no effect of initial depth on escape responses. In conclusion, it appears that the main benefit of increased group density in O. septentrionalis tadpoles is likely to be predator dilution, and that variation in densities of tadpoles influences the escape behaviour of individual tadpoles, regardless of tadpole size.

The power of the seasons: rainfall triggers parental care in poison frogs

The quality of breeding sites is of great importance for the reproductive success and accordingly the fitness of many animal species. Hence, individuals should decide carefully where to rear their offspring. Often parents have to account for multiple characteristics of habitat quality at once, which in turn might change over time. Specimens confronted with such variability may evolve the ability to display context-dependant decision plasticity. Anuran amphibians breeding in ephemeral pools largely face two risks for their offspring: desiccation and predation. The Neotropical poison frog Ranitomeya variabilis deposits both eggs and tadpoles in phytotelmata. These small tadpole nurseries lower the risk of offspring predation. However, because most poison frog tadpoles are cannibalistic, even these pools need to be surveyed for predators, and parents tend to avoid deposition with conspecifics. We tested if this avoidance behaviour does change in parental R. variabilis depending on seasonal circumstances. Over several months we provided the frogs the option to deposit their eggs or tadpoles in pools that did and did not contain chemical cues of cannibalistic conspecifics, respectively. During the rainy season, frogs strongly avoided conspecific cues for both eggs and tadpoles. Anyway, with the change to the dry season, parental preferences changed such that parent frogs were more likely to deposit tadpoles (but not eggs) in pools containing cues of conspecific tadpoles. We suggest that R. variabilis, a species that typically isolates its cannibalistic offspring, has evolved a plastic feeding behaviour with regard to the risk of phytotelmata desiccation. We interpret that parents provide older tadpoles with younger ‘trophic’ tadpoles in order to accelerate their development and save them from impending desiccation.

Making Friends: Social Attraction in Larval Green and Golden Bell Frogs, Litoria aurea

Socio-ecological models combine environmental and social factors to explain the formation of animal groups. In anurans, tadpole aggregations have been reported in numerous species, but the factors driving this behaviour remain unclear. We conducted controlled choice experiments in the lab to determine whether green and golden bell frog (Litoria aurea) tadpoles are directly attracted to conspecifics (social factors) in the absence of environmental cues. Using repeated measures, we found that individual tadpoles strongly preferred associating with conspecifics compared to being alone. Furthermore, this preference was body size dependent, and associating tadpoles were significantly smaller than non-associating tadpoles. We suggest that small tadpoles are more vulnerable to predation and therefore more likely to form aggregations as an anti-predator behaviour. We demonstrate that tadpoles present an ideal model system for investigating how social and ecological factors influence group formation in vertebrates.

Effects of size and size structure on predation and inter-cohort competition in red-eyed treefrog tadpoles

Individual and relative body size are key determinants of ecological performance, shaping the strength and types of interactions within and among species. Size-dependent performance is particularly important for iteroparous species with overlapping cohorts, determining the ability of new cohorts to invade habitats with older, larger conspecifics. We conducted two mesocosm experiments to examine the role of size and size structure in shaping growth and survival in tadpoles of the red-eyed treefrog (Agalychnis callidryas), a tropical species with a prolonged breeding season. First, we used a response surface design to quantify the competitive effect and response of two tadpole size classes across three competitive environments. Large tadpoles were superior per capita effect competitors, increasing the size difference between cohorts through time at high resource availability. Hatchlings were better per biomass response competitors, and maintained the size difference between cohorts when resource availability was low. However, in contrast to previous studies, small tadpoles never closed the size gap with large tadpoles. Second, we examine the relationship between body size, size structure, and predation by dragonfly nymphs (Anax amazili) on tadpole survival and growth. Hatchlings were more vulnerable to predation; predator and large competitor presence interacted to reduce hatchling growth. Again, the size gap between cohorts increased over time, but increased marginally more with predators present. These findings have implications for understanding how variation in resources and predation over the breeding season will shape population size structure through time and the ability of new cohorts to invade habitats with older conspecifics.