Thermal and energetic consequences of nest location and breeding times in Water Pipits (Anthus spinoletta)

Summary The thermal environment has pronounced effects on the energy costs of thermoregulation and affects an animal's allocation of energy to self-maintenance and parental care. Consequently, the selection of reproductive periods, breeding habitats and nest-sites with a favourable microclimate can be advantageous, especially for birds breeding in harsh environments. In this study on Alpine Water Pipits (Anthus spinoletta), we evaluate the importance of spatial and temporal factors on thermoregulatory costs by combining laboratory measurements of metabolic rates under various temperatures with standard operative temperatures (T_es) recorded in the field in different microhabitats. Using these measurements we estimate the thermal and energetic consequences of nest locality and timing of reproduction. Our results show: (1) In the morning, T_es values were much higher on the east-north-east (ENE) slope of a valley than on the west-south-west (WSW) slope; in the afternoon this pattern was reversed. As a consequence, energy costs (E_hour) for thermoregulation on the ENE slope were up to 0.6 RMR (resting metabolic rate at night) lower than on the WSW slope during morning hours and about 0.8 RMR higher during afternoon hours. (2) During the incubation and nestling phases of first and second broods, total energy expenditure for thermoregulation in the daytime (E_daytime) was 0.2–0.3 RMR higher on the ENE slope than on the WSW slope. (3) Within slopes, E_daytime was lower during second broods than during first broods, with differences of 0.06–0.07 RMR during incubation and of 0.32 RMR during nestling care. These differences correspond to the flying costs of females incubating eggs (0.09 RMR) and rearing nestlings (0.25 RMR). We conclude that nest placement in relation to microclimate can improve the female's energy budget, both in terms of the total daily expenditure and its diurnal pattern. From thermal considerations alone, delaying breeding into mid-summer would be advantageous, but this advantage is probably outweighed by the reduced chances for second and replacement clutches and by the necessity to complete moult before migration.

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Thermische und energetische Konsequenzen von Brutplätzen und Brutzeiten bei Wasserpiepern (Anthus spinoletta)

Zusammenfassung Die Umgebungstemperatur beeinflusst den Energieaufwand für die Thermoregulation und bestimmt somit, wie viel Energie ein Tier für die Selbsterhaltung benötigt und wie viel es für Brutpflege aufwenden kann. Daher ist es vorteilhaft, für die Fortpflanzung Zeiten und Orte mit besonders günstigen mikroklimatischen Bedingungen zu wählen. Das gilt insbesondere für Vögel, die in unwirtlichen Biotopen wie z. B. alpinen Regionen brüten. Wir untersuchten den Einfluss von Brutzeit und -ort auf die energetischen Kosten der Thermoregulation am Beispiel einer Population des Wasserpiepers (Anthus spinoletta) im Dischmatal bei Davos (Schweiz). Dazu verknüpften wir drei verschiedene Datensätze: a) Ruhestoffwechsel (RMR), gemessen an gefangenen Vögeln unter verschiedenen Temperaturen, b) Umgebungs- und Nesttemperaturen im Freiland während der Brutzeit und c) standardisierte operative Temperaturen (T_es), ermittelt mit Kupfervögeln in verschiedenen Mikrohabitaten.Die Ergebnisse zeigen: (1) In den Morgenstunden sind T_es-Werte am Ost-Nord-Ost-Hang (ONO) deutlich höher als am West-Süd-West-Hang (WSW); am Nachmittag sind die Verhältnisse umgekehrt. Als Folge davon sind die Energiekosten (E_hour) der Thermoregulation am ONO-Hang in den Morgenstunden bis zu 0.6 RMR niedriger und nachmittags ca. 0.8 RMR höher als am WSW-Hang. (2) In der Bebrütungs- und Nestlingsphase von Erst-und Zweitbruten war der gesamte Energieaufwand für Thermoregulation während der Tagesstunden (E_daytime) am ONO-Hang 0.2–0.3 RMR höher als am WSW-Hang. (3) Innerhalb jedes Hanges war E_daytime bei Zweitbruten niedriger als bei Erstbruten: 0.06–0.07 RMR während der Bebrütung der Eier und 0.32 RMR während der Nestlingsphase. Diese Größenordnungen entsprechen den Flugkosten von Weibchen während der Bebrütungszeit (0.09 RMR) bzw. der Jungenaufzucht (0.25 RMR).Unsere Ergebnisse belegen, dass die Wahl von klimatisch günstigen Habitaten die Energiebilanz von Brutvögeln verbessern kann, sowohl in Hinblick auf den täglichen Gesamtaufwand als auch bezüglich der Verteilung des Energieverbrauchs über den Tag. Aus thermoregulatorischen Gründen wäre auch eine zeitliche Verlagerung des Brutgeschäfts in die Sommermonate (Juli, August) vorteilhaft. Das würde jedoch die Möglichkeiten für Ersatz-und Zweitbruten verringern und die Zeit für die Mauser vor dem Herbstzug verkürzen. Da in unserem Untersuchungsgebiet diese Faktoren für eine erfolgreiche Fortpflanzung eine größere Bedeutung haben als der Energieaufwand für die Thermoregulation, ist der frühe — energetisch 6–32 % teurere — Brutbeginn der Vögel sinnvoll.

     

Tackling unresolved questions in forest ecology: The past and future role of simulation models

1. Understanding the processes that shape forest functioning, structure, and diversity remains challenging, although data on forest systems are being collected at a rapid pace and across scales. Forest models have a long history in bridging data with ecological knowledge and can simulate forest dynamics over spatio‐temporal scales unreachable by most empirical investigations.
2. We describe the development that different forest modelling communities have followed to underpin the leverage that simulation models offer for advancing our understanding of forest ecosystems.
3. Using three widely applied but contrasting approaches – species distribution models, individual‐based forest models, and dynamic global vegetation models – as examples, we show how scientific and technical advances have led models to transgress their initial objectives and limitations. We provide an overview of recent model applications on current important ecological topics and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.
4. Synthesis. This overview shows that forest models, due to their complementarity and mutual enrichment, represent an invaluable toolkit to address a wide range of fundamental and applied ecological questions, hence fostering a deeper understanding of forest dynamics in the context of global change.

Forest models can help understanding the processes that shape forest functioning, structure and diversity, since they can can simulate forest dynamics over spatio‐temporal scales unreachable by most empirical investigations. Here we describe the development of three widely applied but contrasting forest mo−delling approaches — species distribution models, individual‐based models and dynamic global vegetation models. We provide an overview of recent model applications and pinpoint ten key questions that could, and should, be tackled with forest models in the next decade.     

Two-Point Microrheology of Phase-Separated Domains in Lipid Bilayers

Though the importance of membrane fluidity for cellular function has been well established for decades, methods for measuring lipid bilayer viscosity remain challenging to devise and implement. Recently, approaches based on characterizing the Brownian dynamics of individual tracers such as colloidal particles or lipid domains have provided insights into bilayer viscosity. For fluids in general, however, methods based on single-particle trajectories provide a limited view of hydrodynamic response. The technique of two-point microrheology, in which correlations between the Brownian dynamics of pairs of tracers report on the properties of the intervening medium, characterizes viscosity at length-scales that are larger than that of individual tracers and has less sensitivity to tracer-induced distortions, but has never been applied to lipid membranes. We present, to our knowledge, the first two-point microrheological study of lipid bilayers, examining the correlated motion of domains in phase-separated lipid vesicles and comparing one- and two-point results. We measure two-point correlation functions in excellent agreement with the forms predicted by two-dimensional hydrodynamic models, analysis of which reveals a viscosity intermediate between those of the two lipid phases, indicative of global fluid properties rather than the viscosity of the local neighborhood of the tracer.     

Variation in Sex Ratio and Evolutionary Rate of Hemiclonal Rana esculenta Populations

In many plant and animal taxa mutation rates are higher in males than in females. As a result, the evolutionary speed of genes depends on how much time they spend in either sex. Usually, this time differs between genes located on sex chromosomes but not between those on autosomes. Here we present an unusual system with a partially sex-linked inheritance of autosomes: the hemiclonal frog Rana esculenta (E) which is originally a hybrid between the sexual species R. lessonae (L) and R. ridibunda (R). Rana esculenta excludes the L genome prior to meiosis, produces eggs or sperm containing an unrecombined R genome and restores hybridity by mating with R. lessonae (‘hybridogenesis’). Matings between L males and E females result in offspring with an even sex ratio, whereas the reverse combination produces only daughters. The extent of the resulting female bias and the proportion that R alleles have spent in either sex depend on the relative survival (b) and the relative reproductive contribution (a) of E males vs. E females. In this paper, we analyze mathematically how different combinations of a and b influence the sex ratio in R. esculenta populations and, combined with the male/female mutation rate ratio (α), the evolutionary rate of the clonally transmitted R genome. We find that this rate is higher than in an asexual population and lower than in a sexual one. Hence, clonal diversity through new mutations is more easily achievable than in purely asexual species. In contrast, the occurrence and accumulation of deleterious mutations is lower than in a comparable sexual species. We conclude that these intermediate mutation rates improve the ecological and evolutionary potential of hemiclonal organisms, and we draw attention to the implications for the use of microsatellites. Co-ordinating editor: L. Hurst     

Macrophage mobilization and morphology during lens regeneration from the iris epithelium in newts: studies with correlated scanning and transmission electron microscopy

The lens was removed from both eyes of adult newts (Notophthalmus viridescens), and the eyes were fixed in Karnovsky's fixative every 2 days 0-20 days after operation. Anterior half-eyes were prepared by standard procedures for scanning electron microscopy of the surface. Before fixation, the posterior iris surface was cleaned of adhering vitreous mechanically with forceps or by treatment with bovine testicular hyaluronidase or with hyaluronidase and collagenase. Some specimens were cryofractured in buffer or ethanol transverse to the mid-dorsal iris, and the fractured surface viewed with scanning electron microscopy (SEM). Cells with various combinations of ridges, blebs, filopodia, and lamellipodia were observed adhering to the posterior surface of the iris by 6 days after lentectomy. These cells, which exhibited the surface characteristics of macrophages, became more numerous in specimens fixed after longer intervals. Invasion of the iris epithelium was observed in a cryofractured specimen. After observations with SEM, selected specimens were embedded in plastic and sectioned for study with transmission electron microscopy (TEM). The cells on the iris surface had the cytological characteristics of macrophages, and other macrophages were located within the iris epithelium. In specimens fixed 16 or more days after lentectomy, a bulging lens vesicle was regenerating from the dorsal pupillary margin of the iris. Macrophages were absent or few on the surface of this developing lens but remained scattered over the adjoining iris. Roles that might be played by these macrophages during the transdifferentiation of iris epithelium into lens are discussed.