A universal power law for modelling the growth and form of teeth, claws, horns, thorns, beaks, and shells.

BACKGROUND: A major goal of evolutionary developmental biology is to discover general models and mechanisms that create the phenotypes of organisms. However, universal models of such fundamental growth and form are rare, presumably due to the limited number of physical laws and biological processes that influence growth. One such model is the logarithmic spiral, which has been purported to explain the growth of biological structures such as teeth, claws, horns, and beaks. However, the logarithmic spiral only describes the path of the structure through space, and cannot generate these shapes. RESULTS: Here we show a new universal model based on a power law between the radius of the structure and its length, which generates a shape called a 'power cone'. We describe the underlying 'power cascade' model that explains the extreme diversity of tooth shapes in vertebrates, including humans, mammoths, sabre-toothed cats, tyrannosaurs and giant megalodon sharks. This model can be used to predict the age of mammals with ever-growing teeth, including elephants and rodents. We view this as the third general model of tooth development, along with the patterning cascade model for cusp number and spacing, and the inhibitory cascade model that predicts relative tooth size. Beyond the dentition, this new model also describes the growth of claws, horns, antlers and beaks of vertebrates, as well as the fangs and shells of invertebrates, and thorns and prickles of plants. CONCLUSIONS: The power cone is generated when the radial power growth rate is unequal to the length power growth rate. The power cascade model operates independently of the logarithmic spiral and is present throughout diverse biological systems. The power cascade provides a mechanistic basis for the generation of these pointed structures across the tree of life.

Triad hybridization via a conduit species.

Introgressive hybridization can affect the evolution of populations in several important ways. It may retard or reverse divergence of species, enable the development of novel traits, enhance the potential for future evolution by elevating levels of standing variation, create new species, and alleviate inbreeding depression in small populations. Most of what is known of contemporary hybridization in nature comes from the study of pairs of species, either coexisting in the same habitat or distributed parapatrically and separated by a hybrid zone. More rarely, three species form an interbreeding complex (triad), reported in vertebrates, insects, and plants. Often, one species acts as a genetic link or conduit for the passage of genes (alleles) between two others that rarely, if ever, hybridize. Demographic and genetic consequences are unknown. Here we report results of a long-term study of interbreeding Darwin's finches on Daphne Major island, Galápagos. Geospiza fortis acted as a conduit for the passage of genes between two others that have never been observed to interbreed on Daphne: Geospiza fuliginosa, a rare immigrant, and Geospiza scandens, a resident. Microsatellite gene flow from G. fortis into G. scandens increased in frequency during 30 y of favorable ecological conditions, resulting in genetic and morphological convergence. G. fortis, G. scandens, and the derived dihybrids and trihybrids experienced approximately equal fitness. Especially relevant to young adaptive radiations, where species differ principally in ecology and behavior, these findings illustrate how new combinations of genes created by hybridization among three species can enhance the potential for evolutionary change.

The sandwich structure of keratinous layers controls the form and growth orientation of chicken rhinotheca.

The upper beak bone of birds is known to be overlain by the rhinotheca, which is composed of the horny sheath of keratinous layers. However, the details of the structure and growth pattern of the rhinotheca are yet to be understood. In this study, the microstructure of the rhinotheca from chicken specimens of different growth stages (ranging from 1 to ~ 80 days old) was analyzed using a combination of thin section and scanning electron microscopy observations, and small-angle X-ray scattering analysis. We found that the rhinotheca comprises three different layers - outer, intermediate, and inner layers - throughout its growth. The outer layer arises from the proximal portion of the beak bone and covers the dorsal surface of the rhinotheca, whereas the intermediate and inner layers originate in the distal portion of the beak bone and underlie the outer layer. This tri-layered structure of the rhinotheca was also observed in wild bird specimens (grey wagtail, king quail, and brown dipper). On the median plane, micro-layers making up the outer and inner layers are bedded nearly parallel to the rostral bone at the base. However, more distally positioned micro-layers of the outer layer are more anteverted distally. The micro-layers of the intermediate layer are bedded nearly perpendicular to those of the outer and inner layers on the median plane. The growth of micro-layers in the intermediate layer adds thickness to the rhinotheca, which causes the difference in profile between the beak bone and the rhinotheca in the distal portion of the beak. Moreover, the entire intermediate layer grows distally as new proximal micro-layers form. The outer layer is dragged distally by the intermediate layer as a result of its distal growth, for the three layers are closely packed to each other at their boundaries. Furthermore, the occurrence of the intermediate and inner layers in the distal portion of the rostral bone may be because the distal end of the beak is frequently used and worn, and the rhinotheca therefore needs to be replaced more frequently at the distal end. The rhinotheca structure described here will be an important and useful factor in the reconstruction of the beaks of birds in extinct taxa.

Osteogenic Evaluation of Goose-beak Bones According to Processing Temperature.

BACKGROUND/AIM: As an alternative material to autogenous bone, goose-beak bone particles (GBP) have been attracting great attention as a bone substitute due to their biological properties. This study was performed to assess bone generation using GBP in calvarial defects in a rat model. The study focus was the osteogenic potential of goose-beak bone at different processing temperatures. MATERIALS AND METHODS: There were three experimental groups: Control group (critical defect only), low-temperature (LT) group (filled with GBP heat-treated for 20 h at 400°C), and high-temperature (HT) group (filled with GBP heat-treated for 3 h at 1,200°C). RESULTS: The Ca/P atomic ratio of the goose-beak bone was 1.63, and the bones had a bony structure with open pores and interconnected rod-like struts. Micro-computed tomographic analysis revealed the quantity of new bone formation of the HT group was higher than that of the LT group. At 12 weeks after GBP insertion, new bone formation was significantly higher (p<0.05) in the HT group, and there were more new osteocytes in the lacuna in the HT group than in the LT group. Thus, GBP treated at a high temperature formed more new bone than that treated at a low temperature. CONCLUSION: It can be concluded that HT-treated GBP is a graft material that can be effective in promoting bone formation.

Oviposition drives hatching order and developmental disparities with brood mates.

Birth or hatching order can affect fitness. It has long been assumed that the order in which a bird's egg is laid (its oviposition) and first exposed to incubation relative to other eggs in the nest determines the order in which it hatches and the subsequent effects on development and survival. To my knowledge, this cause of hatching order has not been tested while controlling for laying-order effects on egg composition. Like those of many species, the last-laid eggs in clutches of the Lincoln's sparrow ( Melospiza lincolnii) hatch last and produce nestlings with reduced growth and survival rates. Last-hatched Lincoln's sparrows also develop wider bills that are associated with reduced attractiveness of adult courtship songs. Using wild, free-ranging Lincoln's sparrows, I replaced the first or fourth freshly laid egg in four-egg host nests with a freshly laid first egg from a donor nest. I discovered that the experimental elevation of oviposition-which controlled for egg composition-delayed hatching, inhibited growth and widened bill gape, thus confirming the mechanism for hatching order and revealing a potential role for oviposition in reproductive success. Similar effects of postnatal family position could affect fitness in other taxa, as well.

Spatially restricted dental regeneration drives pufferfish beak development.

Vertebrate dentitions are extraordinarily diverse in both morphology and regenerative capacity. The teleost order Tetraodontiformes exhibits an exceptional array of novel dental morphologies, epitomized by constrained beak-like dentitions in several families, i.e., porcupinefishes, three-toothed pufferfishes, ocean sunfishes, and pufferfishes. Modification of tooth replacement within these groups leads to the progressive accumulation of tooth generations, underlying the structure of their beaks. We focus on the dentition of the pufferfish (Tetraodontidae) because of its distinct dental morphology. This complex dentition develops as a result of (i) a reduction in the number of tooth positions from seven to one per quadrant during the transition from first to second tooth generations and (ii) a dramatic shift in tooth morphogenesis following the development of the first-generation teeth, leading to the elongation of dental units along the jaw. Gene expression and 1,1'-Dioctadecyl-3,3,3',3'-tetramethylindocarbocyanine perchlorate (DiI) lineage tracing reveal a putative dental epithelial progenitor niche, suggesting a highly conserved mechanism for tooth regeneration despite the development of a unique dentition. MicroCT analysis reveals restricted labial openings in the beak, through which the dental epithelium (lamina) invades the cavity of the highly mineralized beak. Reduction in the number of replacement tooth positions coincides with the development of only four labial openings in the pufferfish beak, restricting connection of the oral epithelium to the dental cavity. Our data suggest the spatial restriction of dental regeneration, coupled with the unique extension of the replacement dental units throughout the jaw, are primary contributors to the evolution and development of this unique beak-like dentition.

Structure and growth pattern of the bizarre hemispheric prominence on the rostrum of the fossil beaked whale Globicetus hiberus (Mammalia, Cetacea, Ziphiidae).

The rostrum of most ziphiids (beaked whales) displays bizarre swollen regions, accompanied with extreme hypermineralisation and an alteration of the collagenous mesh of the bone. The functional significance of this specialization remains obscure. With the voluminous and dense hemispheric excrescence protruding from the premaxillae, the recently described fossil ziphiid Globicetus hiberus is the most spectacular case. This study describes the histological structure and interprets the growth pattern of this unique feature. Histologically, the prominence in Globicetus is made up of an atypical fibro-lamellar complex displaying an irregular laminar organization and extreme compactness (osteosclerosis). Its development is suggested to have resulted from a protraction of periosteal accretion over the premaxillae, long after the end of somatic growth. Complex shifts in the geometry of this tissue are likely to have occurred during its accretion and no indication of Haversian remodeling could be found. X-ray diffraction and Raman spectroscopy indicate that the bone matrix in the premaxillary prominence of Globicetus closely resembles that of the rostrum of the extant beaked whale Mesoplodon densirostris: apatite crystals are of common size and strongly oriented, but the collagenous meshwork within bone matrix seems to be extremely sparse. These morphological and structural data are discussed in the light of functional interpretations proposed for the highly unusual and diverse ziphiid rostrum. J. Morphol. 277:1292-1308, 2016. © 2016 Wiley Periodicals, Inc.

Microscopic and immunohistochemical study on the cornification of the developing beak in the turtle Emydura macquarii.

The development and cornification of the ramphoteca (beak) in turtles are not known. The microscopic aspects of beak formation have been analyzed in the pleurodirian turtle Emydura macquarii using histological, immunocytochemical and ultrastructural methods. At embryonic Stage 15 the maxillar beak is originated from discontinuous placodes (one frontal and two oral) formed in the epidermis above and below the mouth that later merge into the epidermis of the beak. The mandibular beak is formed by two lateral placodes. In the placodes, basal keratinocytes in contact with local mesenchymal condensations become columnar, and generate suprabasal cells forming 5-6 layers of embryonic epidermis at Stages 17-20 and a compact shedding alpha-layer at the base of the embryonic epidermis. These keratinocytes contain irregular or aggregated reticular bodies made of 30-40 nm thick strands of coarse filaments, mixed with tonofilaments and sparse lipid droplets. Beneath the shedding layer are present 3-4 layers of keratinocytes accumulating coarse filaments mixed with beta-corneous packets, and underneath spindle-shaped beta-cells differentiate where beta-corneous packets completely replace the reticulate bodies. Differently from scales where corneocytes partially merge, beak corneocytes remain separated but they are joined by numerous interlocking spines. The production of beta-cells in the thick corneous layer of the developing beak, like in claws, occurs before the differentiation of beta-cells in the body scutes. This indicates that a massive mesenchymal condensation triggers beta-differentiation before this process is later activated in most of body scutes of the carapace and plastron. J. Morphol. 277:1309-1319, 2016. © 2016 Wiley Periodicals, Inc.

Avian malaria in a boreal resident species: long-term temporal variability, and increased prevalence in birds with avian keratin disorder.

The prevalence of vector-borne parasitic diseases is widely influenced by biological and ecological factors. Environmental conditions such as temperature and precipitation can have a marked effect on haemosporidian parasites (Plasmodium spp.) that cause malaria and those that cause other malaria-like diseases in birds. However, there have been few long-term studies monitoring haemosporidian infections in birds in northern latitudes, where weather conditions can be highly variable and the effects of climate change are becoming more pronounced. We used molecular methods to screen more than 2,000 blood samples collected from black-capped chickadees (Poecile atricapillus), a resident passerine bird. Samples were collected over a 10 year period, mostly during the non-breeding season, at seven sites in Alaska, USA. We tested for associations between Plasmodium prevalence and local environmental conditions including temperature, precipitation, site, year and season. We also evaluated the relationship between parasite prevalence and individual host factors of age, sex and presence or absence of avian keratin disorder. This disease, which causes accelerated keratin growth in the beak, provided a natural study system in which to test the interaction between disease state and malaria prevalence. Prevalence of Plasmodium infection varied by year, site, age and individual disease status but there was no support for an effect of sex or seasonal period. Significantly, birds with avian keratin disorder were 2.6 times more likely to be infected by Plasmodium than birds without the disorder. Interannual variation in the prevalence of Plasmodium infection at different sites was positively correlated with summer temperatures at the local but not statewide scale. Sequence analysis of the parasite cytochrome b gene revealed a single Plasmodium spp. lineage, P43. Our results demonstrate associations between prevalence of avian malaria and a variety of biological and ecological factors. These results also provide important baseline data that will be informative for predicting future changes in Plasmodium prevalence in the subarctic.

Characterization of hatch-size and growth rates of captive and wild-reared brown kiwi (Apteryx mantelli) chicks.

Avian growth rate patterns represent a trade off between a tissue's functional maturity and its capacity for growth. At the time of hatch, the brown kiwi (Apteryx mantelli) limb has a high level of maturity in order for the chick to be able to kick its way out of the shell and walk and forage independently from an early age. Growth curves of limb segments, bill length and bodyweight are presented for captive-reared, BNZ Operation Nest Egg™ chicks over a period of 3 months from the point of hatch. Some parameters were slightly larger in the females than in males at time of hatch, including the bill length. Growth in bodyweight began to slow earlier in males than in females. Regressions of limb and bill measurements over time showed linear patterns of growth instead of a sigmoidal curve as seen in other birds, probably due to the short period of observation. Bodyweight and bill length were then compared to these morphometrics in a wild population of kiwi. Captive-reared chicks were found to hatch with shorter bills and to increase in bodyweight at a faster rate than the wild birds. Rapid weight gain has been implicated in developmental limb deformities in other precocial and long-legged birds and should be avoided in captive kiwi.

Post-hatch heat warms adult beaks: irreversible physiological plasticity in Japanese quail.

Across taxa, the early rearing environment contributes to adult morphological and physiological variation. For example, in birds, environmental temperature plays a key role in shaping bill size and clinal trends across latitudinal/thermal gradients. Such patterns support the role of the bill as a thermal window and in thermal balance. It remains unknown whether bill size and thermal function are reversibly plastic. We raised Japanese quail in warm (30°C) or cold (15°C) environments and then at a common intermediate temperature. We predicted that birds raised in cold temperatures would develop smaller bills than warm-reared individuals, and that regulation of blood flow to the bill in response to changing temperatures would parallel the bill's role in thermal balance. Cold-reared birds developed shorter bills, although bill size exhibited 'catch-up' growth once adults were placed at a common temperature. Despite having lived in a common thermal environment as adults, individuals that were initially reared in the warmth had higher bill surface temperatures than cold-reared individuals, particularly under cold conditions. This suggests that blood vessel density and/or the control over blood flow in the bill retained a memory of early thermal ontogeny. We conclude that post-hatch temperature reversibly affects adult bill morphology but irreversibly influences the thermal physiological role of bills and may play an underappreciated role in avian energetics.

Seasonal variation in sensitivity of the photoperiodic response system in the subtropical tree sparrow (Passer montanus).

We investigated seasonal variations in response to photoperiods in subtropical tree sparrow (Passer montanus), by examining the effects of long and short days on stimulation of response and termination of refractoriness, respectively. We also assessed whether photoperiodic effects were sex dependent. In one study, wild sparrows were transferred in each month of the year to artificial long days (14L/10D) for 12 weeks. Birds transferred from November to March (female) or April (male) showed gradual increase in gonadal growth and darkening of bill color, while those transferred from April (female) or May (male) to July underwent gradual regression in the above responses. Moult in the wing primaries and body feathers progressed with gonadal regression and the birds transferred from April/May to October exhibited gradual increase and decrease in feathers moult. In another study, 6 weeks of short day (9L/15D) exposure could recover responsivity to long days in refractory birds. Male and female birds responded almost in similar fashion with some variations. Overall, our results show a gradual change over seasons in responsiveness of the endogenous response system to stimulatory effects of long day length. They suggest roles of both long and short day lengths in regulation of seasonal cycles in subtropical tree sparrows.

Evidence of accelerated beak growth associated with avian keratin disorder in black-capped chickadees (Poecile atricapillus).

We recently documented an epizootic of beak deformities in more than 2,000 Blackcapped Chickadees (Poecile atricapillus) and other wild bird species in North America. This emerging avian disease, which has been termed avian keratin disorder, results in gross overgrowth of the rhamphotheca, the outer, keratinized layer of the beak. To test the hypothesis that the beak deformities characteristic of this disorder are associated with accelerated keratin production, we measured rates of beak growth and wear in affected Black-capped Chickadees (n=16) and a control sample of unaffected chickadees (n=14) collected from south-central (61°09'-61°38'N, 149°11' -149°48'W) and interior Alaska (64°51' -64°53'N, 147°49' -147°59'W). Rates of absolute growth were 50-100% higher in affected birds than they were in control birds and exceeded records from other passerine species. These results suggest that abnormally rapid epidermal growth is the primary physical mechanism by which beak deformities develop and are maintained in affected chickadees. Although beak overgrowth typically worsened over time, differential patterns of wear influenced the severity and morphology of deformities. In some cases, the effects of accelerated keratin growth were partially mitigated by frequent breakage of rhamphothecal tips. However, mortalities occurred in 9 of 16 birds (56%) with beak deformities during the study, suggesting that avian keratin disorder results in severe health consequences for affected birds. Additional study of factors that control beak keratin production is needed to understand the pathogenesis of this debilitating disease in wild birds.

Growth and development of chicks of two species of partridge: the grey partridge (Perdix perdix) and the chukar (Alectoris chukar).

1. In two partridge species, the grey partridge (Perdix perdix) and chukar (Alectoris chukar), from hatching up to 120 d, the growth rate and development of body mass, wing, tarsus, and bill length were measured and fitted by Gompertz equations. 2. As a typical precocial species, partridges hatched with relatively well developed legs and bills, and wing growth followed a gradual development of thermoregulation. 3. Gompertz growth constants for body mass growth were 0·039 and 0·038 for grey partridges and chukars, respectively. 4. The allometric relationship between tarsus length and body mass followed a geometric similarity (1/3 power) in both grey partridges and chukars.

Distinct development of peripheral trigeminal pathways in the platypus (Ornithorhynchus anatinus) and short-beaked echidna (Tachyglossus aculeatus).

The extant monotremes (platypus and echidnas) are believed to all be capable of electroreception in the trigeminal pathways, although they differ significantly in the number and distribution of electroreceptors. It has been argued by some authors that electroreception was first developed in an aquatic environment and that echidnas are descended from a platypus-like ancestor that invaded an available terrestrial habitat. If this were the case, one would expect the developmental trajectories of the trigeminal pathways to be similar in the early stages of platypus and short-beaked echidna development, with structural divergence occurring later. We examined the development of the peripheral trigeminal pathway from snout skin to trigeminal ganglion in sectioned material in the Hill and Hubrecht collections to test for similarities and differences between the two during the development from egg to adulthood. Each monotreme showed a characteristic and different pattern of distribution of developing epidermal sensory gland specializations (electroreceptor primordia) from the time of hatching. The cross-sectional areas of the trigeminal divisions and the volume of the trigeminal ganglion itself were also very different between the two species at embryonic ages, and remained consistently different throughout post-hatching development. Our findings indicate that the trigeminal pathways in the short-beaked echidna and the platypus follow very different developmental trajectories from the earliest ages. These findings are more consistent with the notion that the platypus and echidna have both diverged from an ancestor with rudimentary electroreception and/or trigeminal specialization, rather than the contention that the echidna is derived from a platypus-like ancestor.

[Postnatal growth patterns in eight species of herons and egrets (Ciconiiformes: Ardeidae)]. / Patrones de crecimiento postnatal en ocho especies de garzas (Ciconiiformes: Ardeidae).

Avian postnatal growth has received considerable attention and its ecological implications have been deeply analyzed. In this current paper, I describe the patterns of culmen and tarsus growth, as well as of weight gain patterns in eight species of herons and egrets (Aves: Ardeidae) found in the Birama Swamp in Eastern Cuba. Between 1998 and 2006,714 nestlings of the following species were measured every two days: Butorides virescens, Bubulcus ibis, Egretta thula, E. tricolor, E. caerulea, E. rufescens, Ardea alba and Nycticorax nycticorax. Logistic and Gompertz equations were adjusted to data using non-lineal regression models with adult values as the asymptote. For each species, the following were determined and recorded: growth rate, age at inflexion, instantaneous growth rates at each age interval, and time taken to reach 90% of adult size. Reported hatchling sizes were similar in other localities, with a variation coefficient ranging between 10-19%. At hatch, each species exhibited differing sizes relative to adult values. In all cases, Gompertz equations were best fitted to explain more variance and lesser residuals. Rates of weight change and tarsus growth were alometrically related to the log of adult weight. Two main growth processes were identified: a physical extension in dimensions of each measurement reflecting inter-specific morphometric differences, and a lineal increase of the growth period from Green Heron to Great Egret. The Black-crowned Night Heron, Cattle Egret and Reddish Egret exhibited some unique measurement characteristics in comparison to the remaining members of the family. All results support the hypothesis that hypermorphosis, as the main evolutionary process in the microevolution of Ardeidae, is caused by a delayed final moment of growth.

Patrones de crecimiento postnatal en ocho especies de garzas (Ciconiiformes: Ardeidae)

Postnatal growth patterns in eight species of herons and egrets (Ciconiiformes: Ardeidae). Avian postnatal growth has received considerable attention and its ecological implications have been deeply analyzed. In this current paper, I describe the patterns of culmen and tarsus growth, as well as of weight gain patterns in eight species of herons and egrets (Aves: Ardeidae) found in the Birama Swamp in Eastern Cuba. Between 1998 and 2006, 714 nestlings of the following species were measured every two days: Butorides virescens, Bubulcus ibis, Egretta thula, E. tricolor, E. caerulea, E. rufescens, Ardea alba and Nycticorax nycticorax. Logistic and Gompertz equations were adjusted to data using non-lineal regression models with adult values as the asymptote. For each species, the following were determined and recorded: growth rate, age at inflexion, instantaneous growth rates at each age interval, and time taken to reach 90% of adult size. Reported hatchling sizes were similar in other localities, with a variation coefficient ranging between 10-19%. At hatch, each species exhibited differing sizes relative to adult values. In all cases, Gompertz equations were best fitted to explain more variance and lesser residuals. Rates of weight change and tarsus growth were alometrically related to the log of adult weight. Two main growth processes were identified: a physical extension in dimensions of each measure ment reflecting inter-specific morphometric differences, and a lineal increase of the growth period from Green Heron to Great Egret. The Black-crowned Night Heron, Cattle Egret and Reddish Egret exhibited some unique measurement characteristics in comparison to the remaining members of the family. All results support the hypothesis that hypermorphosis, as the main evolutionary process in the microevolution of Ardeidae, is caused by a delayed final moment of growth. Rev. Biol. Trop. 59 (2): 771-787. Epub 2011 June 01.

El crecimiento en aves forma parte de las estrategias reproductivas y su implicación ecológica ha sido profundamente debatida. En este trabajo se describe el crecimiento del pico, peso y tarso de ocho garzas (Aves: Ardeidae) en la ciénaga de Birama, Cuba. Para ello, entre 1998-2006, se midieron en días alternos un total de 714 pichones de Butorides virescens, Bubulcus ibis, Egretta thula, E. tricolor, E. caerulea, E. rufescens, Ardea alba y Nycticorax nycticorax, durante sus dos primeras semanas de vida. Se hicieron ajustes a las curvas Logística y de Gompertz y se usó regresiones no lineales y valores adultos como asíntotas, además se determinaron las variables que describenel crecimiento. El tamaño de los pichones al eclosionar fue similar al encontrado en otras localidades, con CV entre 10-19% y diferencias relativas al tamaño adulto. Las ecuaciones de Gompertz tuvieron menos residuos en todos los casos. Entonces, se identificaron dos procesos fundamentales de extensión en el crecimiento de cada estructura: uno físico y otro de duración, entre B. virescens y E. alba. Nycticorax, B. ibis y E. rufescens mostraron variaciones que los diferenciaron del resto. Los resultados apoyan la hipótesis de la microevolución de Ardeidae por hipermorfosis, por un retardo en el momento final del crecimiento.

Minor beak trimming in chickens leads to loss of mechanoreception and magnetoreception.

Routine removal of the tip of the beak of chickens within the poultry industry leads to changes in pecking behavior, which have previously been interpreted as being indicative of pain. By analyzing the force of pecks, with and without the topical application of an analgesic to the beak, we investigated if changes in pecking behavior were due to a loss of sensitivity in the beak or were pain related. Pecking behavior was compared between intact-beak and beak-trimmed chicks with or without topical application of lignocaine during a pain-free period (within 24 h of beak trimming) or after this period (d 2 to 9 of age). After pecking behavior tests, chicks were trained to use a magnetic stimulus to locate hidden food in 1 corner of a square arena. In unrewarded magnetic tests, the location of the chick relative to the magnetic stimulus was determined by automatic image recognition. Beak-trimmed chicks pecked harder than intact-beak chicks within 24 h of beak trimming (P = 0.04), possibly as a means of compensating for the loss of sensory feedback in beak-trimmed chicks. At 2 to 9 d of age, beak-trimmed chicks took longer to peck the pecking stimulus (P < 0.001) and showed fewer pecks in total (P < 0.001), suggesting a reduced pecking motivation. The force of pecks, however, did not differ among treatments at 2 to 9 d of age, suggesting that beak-trimmed chicks were not experiencing pain from the beak. In the magnetic tests, hungry intact-beak chicks stayed nearer to the magnetic stimulus (P = 0.005) and spent proportionally more time within 125 mm of the magnetic stimulus (P = 0.02) that had previously been associated with food than beak-trimmed chicks, which indicated that intact-beak birds were better able to detect the magnetic stimulus than beak-trimmed birds. We concluded that minor beak trimming at a young age did not result in pain in young domestic chicks, but instead led to impaired function of the magnetoreceptors and mechanoreceptors of the beak.

Different effects of infrared and one-half hot blade beak trimming on beak topography and growth.

This study examined the effects of infrared beak treatment (IR-BT) and hot blade beak trimming (HB-BT) on beak length and production in laying hen pullets. Seventy-two 1-d-old birds were randomly assigned to the HB-BT, IR-BT, or control group. Birds were pair housed by treatment, and beak images and production indices were obtained posttreatment at fixed intervals for 10 wk. Immediately after treatment, HB-BT beaks were shorter than control or IR-BT beaks (P < 0.01), whereas control and IR-BT beaks remained comparable in length until the onset of tissue degeneration and erosion of the IR-BT beaks at 1 to 2 wk posttreatment. At wk 2 posttreatment, beaks were longest in control birds, intermediate in IR-BT birds, and shortest in HB-BT birds (P < 0.01). Thereafter, an increase in beak length in all treatments was observed over time (P < 0.01), but HB-BT beaks had the greatest regrowth. The beak length of birds in the HB-BT group was similar to that of birds in the IR-BT group from wk 3 to 8, and then grew longer at wk 9 and 10 posttreatment (P < 0.01). The effects of treatments on BW emerged at d 5 posttreatment. The BW of birds in the HB-BT group was suppressed up to and including wk 9 posttreatment compared with that of control birds (P < 0.05) and was significantly lower than that of birds in the IR-BT group between 2 and 4 wk posttreatment (P < 0.05). Birds in the IR-BT group did not differ from control birds after wk 3 posttreatment (P < 0.05). By the final week of the study, differences in BW across treatments were no longer apparent (P > 0.05). For the most part, feed intake was higher in control birds, intermediate in birds in the IR-BT group, and lowest in birds in the HB-BT group until wk 9 posttreatment (P < 0.05). Similarly, feed waste was generally higher in control birds and least in birds in the HB-BT group (P < 0.05). The IR-BT treatment appeared to be more effective at inhibiting beak regrowth, with a less pronounced effect on feed intake than the HB-BT treatment in laying hen pullets.

Scaling and shear transformations capture beak shape variation in Darwin's finches.

Evolution by natural selection has resulted in a remarkable diversity of organism morphologies that has long fascinated scientists and served to establish the first relations among species. Despite the essential role of morphology as a phenotype of species, there is not yet a formal, mathematical scheme to quantify morphological phenotype and relate it to both the genotype and the underlying developmental genetics. Herein we demonstrate that the morphological diversity in the beaks of Darwin's Finches is quantitatively accounted for by the mathematical group of affine transformations. Specifically, we show that all beak shapes of Ground Finches (genus Geospiza) are related by scaling transformations (a subgroup of the affine group), and the same relationship holds true for all the beak shapes of Tree, Cocos, and Warbler Finches (three distinct genera). This analysis shows that the beak shapes within each of these groups differ only by their scales, such as length and depth, which are genetically controlled by Bmp4 and Calmodulin. By measuring Bmp4 expression in the beak primordia of the species in the genus Geospiza, we provide a quantitative map between beak morphology and the expression levels of Bmp4. The complete morphological variation within the beaks of Darwin's finches can be explained by extending the scaling transformations to the entire affine group, by including shear transformations. Altogether our results suggest that the mathematical theory of groups can help decode morphological variation, and points to a potentially hierarchical structure of morphological diversity and the underlying developmental processes.