Differential organ growth during development is essential for adults to maintain the correct proportions and achieve their characteristic shape. Organs scale with body size, a process known as allometry that has been studied extensively in a range of organisms. Such scaling rules, typically studied from a limited sample, are assumed to apply to all members of a population and/or species. Here we study scaling in the compound eyes of workers of the wood ant, Formica rufa, from different colonies within a single population. Workers' eye area increased with body size in all the colonies showing a negative allometry. However, both the slope and intercept of some allometric scaling relationships differed significantly among colonies. Moreover, though mean facet diameter and facet number increased with body size, some colonies primarily increased facet number whereas others increased facet diameter, showing that the cellular level processes underlying organ scaling differed among colonies. Thus, the rules that govern scaling at the organ and cellular levels can differ even within a single population.Understanding how organ size and shape is controlled during development is a major challenge in biology. The control of organ morphology is particularly problematic for organisms that need to develop organs to meet specific requirements under fluctuating conditions and resources. In natural environments, adults from a single species can vary enormously in body size due to a combination of genetic and environmental factors. The changes in organ size that accompany changes in body size can be characterised by allometric scaling relationships. The scaling of any feature with size can be described by:where x is a measure of body size, Y is the size of the organ in question, α is the scaling exponent and b is the initial growth index 1 . When no change occurs in the relative size of an organ with body size (α = 1) the relationship is described as isometric 2 . More typically, however, organs show negative allometries (α < 1) becoming relatively smaller as body size increases 3,4 . Even with negative allometries organs can be absolutely larger in animals with a greater body size, but relatively smaller when compared to smaller conspecifics 1,5,6 . In rare cases, organ size may show positive allometry increasing relatively faster than body size (α > 1) 7,8 . Such positive allometry is often associated with organs under sexual selection 7,8 . The scaling of different organs within a body is the product of differential growth; as an organism grows larger, certain organs grow at a faster rate than others 3,9 . This is thought to occur through differential resource allocation 9,10,11 , whereby resources are distributed to different organs at different rates. Scaling has been studied in many taxa including mammals and birds [12][13][14][15] and especially insects [15][16][17][18][19] . In part, this is due to the power of genetic tools available in the fruit fly Drosophila melanogaster 20 but also because of the mode of development of...

Background and Aims. The presence of novel structures in orchid flowers, including auricles, rostellum and bursicles on the gynostemium and a lobed labellum, has prompted long-standing homology disputes, fuelled by conflicting evidence from a wide range of sources. Re-assessment of this debate using an improved model is timely, following recent phylogenetic insights and on the cusp of a revolution in developmental genetics.Methods. We use new data from floral development and anatomy in the small-flowered terrestrial orchid Herminium monorchis as a model to explore organ homologies in orchid flowers within the context of a review of recent literature on developmental genetics.Key Results. The apex of the median carpel of Herminium is trilobed, and the bursicles develop from its lateral lobes, relatively late in flower ontogeny. The bursicles enclose the viscidia, which adhere to the tapetal remnants to form a caudicle linking the viscidium with the pollinium. The auricles are initiated earlier than the bursicles, but they also remain unvascularized. The deeply trilobed labellum possesses three vascular traces, in contrast with the lateral petals, each of which contains a single vascular trace. The two lateral labellum traces diverge from the traces supplying the two adjacent lateral sepals. Data from flower ontogeny and anatomy conflict with respect to organ homologies.Conclusions. Much progress has recently been made in understanding the exceptional differentiation shown by orchids among perianth segments, focusing on multiple copies of the DEF/AP3 subclass of B-class MADS-box genes. In contrast, untangling homologies of profound congenital union of multiple floral organs forming the orchid gynostemium is hampered by their profound congenital union, which we ascribe to overlap in gene expression between organs. Thus, the functional morphology of the orchid flower could ultimately reflect extreme synorganization and associated genetic integration. Analogizing the deeply lobed orchid labellum with a compound leaf, we speculate that KNOX genes could be implicated not only in their demonstrated role in spur development but also in the development of both the characteristic lobed morphology of the orchid labellum and the lobing of the median carpel that differentiates the bursicles and rostellum.

Environmental and genetic influences cause individuals of a species to differ in size. As they do so, organ size and shape are scaled to available resources whilst maintaining function. The scaling of entire organs has been investigated extensively but scaling within organs remains poorly understood. By making use of the structure of the insect compound eye, we show that different regions of an organ can respond differentially to changes in body size. Wood ant (Formica rufa) compound eyes contain facets of different diameters in different regions. When the animal body size changes, lens diameters from different regions can increase or decrease in size either at the same rate (a ‘grade’ shift) or at different rates (a ‘slope’ shift). These options are not mutually exclusive, and we demonstrate that both types of scaling apply to different regions of the same eye. This demonstrates that different regions within a single organ can use different rules to govern their scaling, responding differently to their developmental environment. Thus, the control of scaling is more nuanced than previously appreciated, diverse responses occurring even among homologous cells within a single organ. Such fine control provides a rich substrate for the diversification of organ morphology.

Artificial lighting at night (ALAN) is increasingly recognised as having negative effects on many organisms, though the exact mechanisms remain unclear. Glow worms are likely susceptible to ALAN because females use bioluminescence to signal to attract males. We quantified the impact of ALAN by comparing the efficacy of traps that mimicked females to attract males in the presence or absence of a white artificial light source (ALS). Illuminated traps attracted fewer males than did traps in the dark. Illuminated traps closer to the ALS attracted fewer males than those further away, whereas traps in the dark attracted similar numbers of males up to 40 m from the ALS. Thus, ALAN impedes females' ability to attract males, the effect increasing with light intensity. Consequently, ALAN potentially affects glow worms' fecundity and long-term population survival. More broadly, this study emphasises the potentially severe deleterious effects of ALAN upon nocturnal insect populations.

Static allometries determine how organ size scales in relation to body mass. The extent to which these allometric relationships are free to evolve, and how they differ among closely related species, has been debated extensively and remains unclear; changes in intercept appear common, but changes in slope are far rarer. Here, we compare the scaling relationships that govern the structure of compound eyes of four closely related ant species from the genus Formica. Comparison among these species revealed changes in intercept but not slope in the allometric scaling relationships governing eye area, facet number, and mean facet diameter. Moreover, the scaling between facet diameter and number was conserved across all four species. In contrast, facet diameters from distinct regions of the compound eye differed in both intercept and slope within a single species and when comparing homologous regions among species. Thus, even when species are conservative in the scaling of whole organs, they can differ substantially in regional scaling within organs. This, at least partly, explains how species can produce organs that adhere to genus wide scaling relationships while still being able to invest differentially in particular regions of organs to produce specific features that match their ecology.

The prevailing account of visually controlled routes is that an ant learns views as it follows a route, while guided by other path-setting mechanisms. Once a set of route views is memorised, the insect follows the route by turning and moving forwards when the view on the retina matches a stored view. We engineered a situation in which this account cannot suffice in order to discover whether there may be additional components to the performance of routes. One-eyed wood ants were trained to navigate a short route in the laboratory, guided by a single black, vertical bar placed in the blinded visual field. Ants thus had to turn away from the route to see the bar. They often turned to look at or beyond the bar and then turned to face in the direction of the goal. Tests in which the bar was shifted to be more peripheral or more frontal than in training produced a corresponding directional change in the ants' paths, demonstrating that they were guided by the bar. Examination of the endpoints of turns towards and away from the bar indicate that ants use the bar for guidance by learning how large a turn-back is needed to face the goal. We suggest that the ants' zigzag paths are in part controlled by turns of a learnt amplitude and that these turns are an integral part of visually guided route following.

Metabolic rate and its relationship with body size is a fundamental determinant of many life history traits and potentially of organismal fitness. Alongside various environmental and physiological factors, the metabolic rate of insects is linked to distinct ventilation patterns. Despite significant attention, however, the precise role of these ventilation patterns remains uncertain. Here, we determined the allometric scaling of metabolic rate and respiratory water loss in the red wood ant, as well as assessing the effect of movement upon metabolic rate and ventilation pattern. Metabolic rate and respiratory water loss are both negatively allometric. We observed both continuous and cyclic ventilation associated with relatively higher and lower metabolic rates, respectively. In wood ants, however, movement not metabolic rate is the primary determinant of which ventilation pattern is performed. Conversely, metabolic rate not ventilation pattern is the primary determinant of respiratory water loss. Our statistical models produced a range of relatively shallow intraspecific scaling exponents between 0.40 and 0.59, emphasising the dependency upon model structure. Previous investigations have revealed substantial variation in morphological allometry among wood ant workers from different nests within a population. Metabolic rate scaling does not exhibit the same variability, suggesting that these two forms of scaling respond to environmental factors in different ways.

Differences in organ scaling among individuals may play an important role in determining behavioural variation. In social insects, there are well-documented intraspecific differences in colony behaviour, but the extent that organ scaling differs within and between colonies remains unclear. Using 12 different colonies of the bumblebee Bombus terrestris , we aim to address this knowledge gap by measuring the scaling relationships between three different organs (compound eyes, wings and antennae) and body size in workers . Though colonies were exposed to different rearing temperatures, this environmental variability did not explain the differences of the scaling relationships. Two colonies had differences in wing versus antenna slopes, three colonies showed differences in wing versus eye slopes and a single colony has differences between eye versus antenna slopes. There are also differences in antennae scaling slopes between three different colonies, and we present evidence for putative trade-offs in morphological investment. We discuss the utility of having variable scaling among colonies and the implication for understanding variability in colony fitness and behaviour.