Tree size and fecundity influence ballistic seed dispersal of two dominant mast-fruiting species in a tropical rain forest

Seed production, seed dispersal, and seedling recruitment are integral to forest dynamics, especially in masting species. Often these are studied separately, yet scarcely ever for species with ballistic dispersal even though this mode of dispersal is common in legume trees of tropical African rain forests. Here, we studied two dominant main-canopy tree species, Microberlinia bisulcata and Tetraberlinia bifoliolata (Caesalpinioideae), in 25 ha of primary rain forest at Korup, Cameroon, during two successive masting events (2007/2010). In the vicinity of c. 100 and 130 trees of each species, 476/580 traps caught dispersed seeds and beneath their crowns c. 57,000 pod valves per species were inspected to estimate tree-level fecundity. Seed production of trees increased non-linearly and asymptotically with increasing stem diameters. It was unequal within the two species’ populations, and differed strongly between years to foster both spatial and temporal patchiness in seed rain. The M. bisulcata trees could begin seeding at 42–44 cm diameter: at a much larger size than could T. bifoliolata (25 cm). Nevertheless, per capita life-time reproductive capacity was c. five times greater in M. bisulcata than T. bifoliolata owing to former’s larger adult stature, lower mortality rate (despite a shorter life-time) and smaller seed mass. The two species displayed strong differences in their dispersal capabilities. Inverse modelling (IM) revealed that dispersal of M. bisulcata was best described by a lognormal kernel. Most seeds landed at 10–15 m from stems, with 1% of them going beyond 80 m (<100 m). The direct estimates of fecundity significantly improved the models fitted. The lognormal also described well the seedling recruitment distribution of this species in 121 ground plots. By contrast, the lower intensity of masting and more limited dispersal of the heavier-seeded T. bifoliolata prevented reliable IM. For this species, seed density as function of distance to traps suggested a maximum dispersal distance of 40–50 m, and a correspondingly more aggregated seedling recruitment pattern ensued than for M. bisulcata. From this integrated field study, we conclude that the reproductive traits of M. bisulcata give it a considerable advantage over T. bifoliolata by better dispersing more seeds per capita to reach more suitable establishment sites, and combined with other key traits they explain its local dominance in the forest. Understanding the linkages between size at onset of maturity, individual fecundity, and dispersal capability can better inform the life-history strategies, and hence management, of co-occurring tree species in tropical forests.

Size does matter: 18 amino acids at the N-terminal tip of an amino acid transporter in Leishmania determine substrate specificity

Long N-terminal tails of amino acid transporters are known to act as sensors of the internal pool of amino acids and as positive regulators of substrate flux rate. In this study we establish that N-termini of amino acid transporters can also determine substrate specificity. We show that due to alternative trans splicing, the human pathogen Leishmania naturally expresses two variants of the proline/alanine transporter, one 18 amino acid shorter than the other. We demonstrate that the longer variant (LdAAP24) translocates both proline and alanine, whereas the shorter variant (∆18LdAAP24) translocates just proline. Remarkably, co-expressing the hydrophilic N-terminal peptide of the long variant with ∆18LdAAP24 was found to recover alanine transport. This restoration of alanine transport could be mediated by a truncated N-terminal tail, though truncations exceeding half of the tail length were no longer functional. Taken together, the data indicate that the first 18 amino acids of the negatively charged N-terminal LdAAP24 tail are required for alanine transport and may facilitate the electrostatic interactions of the entire negatively charged N-terminal tail with the positively charged internal loops in the transmembrane domain, as this mechanism has been shown to underlie regulation of substrate flux rate for other transporters.

Particle concentration and size distribution in the NGRIP ice core

A novel laser microparticle detector used in conjunction with continuous sample melting has provided a more than 1500 m long record of particle concentration and size distribution of the NGRIP ice core, covering continuously the period approximately from 9.5-100 kyr before present; measurements were at 1.65 m depth resolution, corresponding to approximately 35-200 yr. Particle concentration increased by a factor of 100 in the Last Glacial Maximum (LGM) compared to the Preboreal, and sharp variations of concentration occurred synchronously with rapid changes in the delta18O temperature proxy. The lognormal mode µ of the volume distribution shows clear systematic variations with smaller modes during warmer climates and coarser modes during colder periods. We find µ ~ 1.7 µm diameter during LGM and µ ~ 1.3 µm during the Preboreal. On timescales below several 100 years µ and the particle concentration exhibit a certain degree of independence present especially during warm periods, when µ generally is more variable. Using highly simplifying considerations for atmospheric transport and deposition of particles we infer that (1) the observed changes of µ in the ice largely reflect changes in the size of airborne particles above the ice sheet and (2) changes of µ are indicative of changes in long range atmospheric transport time. From the observed size changes we estimate shorter transit times by roughly 25% during LGM compared to the Preboreal. The associated particle concentration increase from more efficient long range transport is estimated to less than one order of magnitude.

Particle size spectrum analysis during Maria S. Merian cruise MSM26, spring 2013

A Laser In-Situ Scattering Transmissometer (LISST) was used to collect vertical distribution data of particles from 2.5 to 500 µm in size. The LISST uses a multi-ring detector to measure scattering light of particles from a laser diode. Particles are classified into 32 log-spaced bins and the concentration of each bin is calculated as micro-liters per liter (µl/l). The instrument is rated to a depth of 300 m, and also records temperature and pressure. The sample interval was set to record every second. The LISST was attached to the LOPC frame to conduct casts and allow for particle-size comparisons between the two instruments. The LOPC is rated to a depth of 2000 m, thus a short deployment to a depth of 300 m was first conducted with both instruments. The instruments were then returned to the deck and the LISST removed via a quick release bracket so deep LOPC casts could be continued at a station. Raw LISST size-spectrum data is presented as concentrations for each of the 32 size bins for every second of the cast.

The relevance of seed size in modulating leaf physiology and early plant performance in two tree species

he size of seeds and the microsite of seed dispersal may affect the early establishment of seedlings through different physiological processes. Here, we examined the effects of seed size and light availability on seedling growth and survival, and whether such effects were mediated by water use efficiency. Acorns of Quercus petraea and the more drought-tolerant Quercus pyrenaica were sowed within and around a tree canopy gap in a sub-Mediterranean forest stand. We monitored seedling emergence and measured predawn leaf water potential (Ψpd), leaf nitrogen per unit area (Na), leaf mass per area, leaf carbon isotope composition (δ13C) and plant growth at the end of the first summer. Survival was measured on the next year. Path analysis revealed a consistent pattern in both species of higher δ13C as Ψpd decreased and higher δ13C as seedlings emerged later in the season, indicating an increase in 13C as the growing season is shorter and drier. There was a direct positive effect of seed size on δ13C in Q. petraea that was absent in Q. pyrenaica. Leaf δ13C had no effect on growth but the probability of surviving until the second year was higher for those seedlings of Q. pyrenaica that had lower δ13C on the first year. In conclusion, leaf δ13C is affected by seed size, seedling emergence time and the availability of light and water, however, leaf δ13C is irrelevant for first year growth, which is directly dependent on the amount of seed reserves.

Regulación del movimiento y anticipación en el resto del primer servicio en tenis

Aunque se han logrado importantes avances en estudios de laboratorio con diseños experimentales poco representativos (e.g., Farrow y Reid, 2012; Nieminen, Piirainen, Salmi, y Linnamo, 2013), a día de hoy, todavía se desconoce a cabalidad cómo los jugadores de tenis de diferente nivel de pericia calibran o ajustan sus movimientos a las demandas espacio-temporales presentes en la tarea de resto de un primer servicio. ! Escasos trabajos se han llevado a cabo in situ y a la mayoría se les puede cuestionar algún aspecto de la metodología empleada. Así pues, en varios estudios la frecuencia de grabación ha sido limitada (e.g., a 50 Hz en Jackson y Gudgeon, 2004; Triolet, Benguigui, Le Runigo y Williams, 2013), o la velocidad del saque ha sido visiblemente inferior a la habitual (cf. Carboch, Süss y Kocib, 2014; Williams, Singer y Weigelt, 1998). También, en algunos estudios los participantes experimentados no han sido jugadores de nivel internacional (e.g., Avilés, Ruiz, Sanz y Navia, 2014), y el tamaño muestral ha sido muy pequeño (e.g., Gillet, Leroy, Thouvarecq, Mégrot y Stein, 2010). ! Además, en los diferentes trabajos se han utilizado una diversidad de métodos e instrumentos de medida y los criterios de codificación del inicio de los movimientos y de las respuestas han diferido; como consecuencia el lapso visomotor de respuesta (LVMr) ha sido muy dispar variando considerablemente de 198 a 410 ms. Considerando los inconvenientes señalados anteriormente, el presente estudio tuvo como objetivo determinar un modelo técnico de regulación temporal de los movimientos y de la respuesta del restador, tomando en cuenta el flujo continuo de información proporcionado por el sacador. Para ello, se realizó un análisis cronométrico de los restos de doce jugadores de diferente nivel deportivo (seis internacionales y seis nacionales) que respondieron de forma natural enviando sus devoluciones hacia las dianas. Se grabaron las acciones de los restadores y sacadores con una cámara Casio Exilim Pro Ex-F1 de alta velocidad (300 Hz) y luego se realizó un análisis imagen por imagen cada 3.33 ms. Una vez obtenidos los datos de los vídeos se realizaron análisis con las pruebas de ANOVA de un factor, ANCOVA con la velocidad del saque como covariable, U de Mann-Whitney y Chi-cuadrado de Pearson. En cuanto a la regulación del movimiento hasta el momento del despegue, los jugadores internacionales iniciaron sus acciones antes que los jugadores nacionales lo que podría indicar una mejor preparación al ejecutar los movimientos como reflejo del nivel de pericia. Los jugadores internacionales iniciaron la elevación del pie posterior a -293 ms y los jugadores nacionales a -202 ms. Todas estas acciones se fueron enlazando unas con otras y fue en el momento del impacto del sacador donde los restadores demostraron una remarcable coordinación perceptivo-motriz. Por consiguiente, los jugadores internacionales despegaron e iniciaron el vuelo a tan solo -6.5 ms del impacto y los jugadores nacionales lo hicieron más tarde a +19.5 ms. A lo largo de la secuencia temporal, todo parece indicar que las informaciones que utilizan los restadores interactúan entre sí; información más temprana y menos fiable para anticipar o moverse antes e información más tardía y más fiable para regular la temporalización de las acciones. Los restadores de nivel internacional y nacional anticiparon a nivel espacial en un bajo porcentaje (7.7% vs. 13.6%) y en tiempos similares (-127 vs. -118 ms) sugiriendo que la utilización de variables ópticas tempranas y menos fiables solo se produce en contadas ocasiones. Por otra parte, estos datos se relacionan con una gran precisión en la respuesta ya que tanto los jugadores internacionales como los nacionales demostraron un alto porcentaje de acierto al responder (95.4% vs. 96.7%). Se había señalado que los jugadores internacionales y nacionales se diferenciarían en el tiempo de caída (i.e., aterrizaje) del primer pie del salto preparatorio, sin embargo ese efecto no fue encontrado (128 vs. 135 ms). Tampoco se hallaron diferencias en el porcentaje de caída con el pie contrario a la dirección de la pelota (58% vs. 62%). Donde sí ambos grupos se diferenciaron fue en el tiempo de caída del segundo pie (147 vs. 168 ms). Esta diferencia de 21 ms fue crucial y fue una prueba de la mayor rapidez de los jugadores internacionales; sugiriendo que ésta acción se podría relacionar con el momento del inicio de la respuesta. Aunque los jugadores internacionales hayan demostrado ser más rápidos en relación con sus capacidades funcionales, ambos grupos no se diferenciaron en todas las variables relacionadas con el LVMr. Ellos no utilizaron esos valiosos milisegundos ganados en el instante de la caída del segundo pie para responder más pronto, ya que el LVMr del miembro superior fue el mismo para ambos grupos (179 vs. 174 ms). Es como si hubiesen tenido todo el tiempo del mundo para seguir ajustando sus acciones hasta el propio golpeo. Además, estos tiempos largos sugieren que en la gran mayoría de los restos la información clave que determinó la respuesta fue detectada (extraída) en momentos cercanos al golpeo del sacador y en la primera parte del vuelo de la pelota. Asimismo, se constató que en general el LVMr se ve influenciado por el tipo de información utilizada. De esta manera, cuando se tomaron en cuenta los ensayos en los que hubo anticipación espacial reflejados en el LVMr del cuerpo entero los tiempos disminuyeron (152 vs. 136 ms). Por otra parte, existieron ocasiones (13%) en los que tanto los jugadores internacionales como los nacionales respondieron tarde recibiendo saques directos (208 vs. 195 ms). Es muy posible que en estos casos los jugadores hayan tenido problemas para detectar la información respondiendo fuera de los márgenes temporales de acción lo que mermó su rendimiento. Lo mismo pudo haber ocurrido cuando ambos grupos de jugadores corrigieron el movimiento del miembro superior tras el impacto (17% vs. 10%) lo que aumentó el tiempo en responder al redirigir la respuesta hacia el lado correcto (208 vs. 205 ms). Además, los jugadores internacionales obtuvieron tiempos de movimiento menores que el de los jugadores nacionales (509 vs. 531 ms) lo que se reflejó en un tiempo total de actuación menor (683 vs. 703 ms). Por último, en cuanto al rendimiento del resto, los jugadores internacionales obtuvieron valores superiores a los jugadores nacionales (1.3 vs. 0.9). ABSTRACT Although there have been significant advances in laboratory studies with unrepresentative experimental designs (e.g., Farrow y Reid, 2012; Nieminen, Piirainen, Salmi, y Linnamo, 2013), today it is still unknown to full extent how tennis players of different levels of expertise calibrate or adjust their movements to the spatial-temporal demands present in the return of a first serve. Few studies have been carried out in situ and some aspects of the methodology most of them used can be questioned. Thus, in several studies the recording frequency has been limited (e.g., a 50 Hz en Jackson y Gudgeon, 2004; Triolet, Benguigui, Le Runigo y Williams, 2013), or serve speed was visibly lower than the usual one (cf. Carboch, Süss y Kocib, 2014; Williams, Singer y Weigelt, 1998). Also, in some studies, experienced participants have not played at international level (e.g., Avilés, Ruiz, Sanz y Navia, 2014), and the sample size has been very small (e.g., Gillet, Leroy, Thouvarecq, Mégrot y Stein, 2010). Furthermore, different works have used a variety of methods and measurement instruments and coding criteria of the onset of movements and responses have differed; due to this, visuomotor response delay (LVMr) has been very uneven, varying considerably from 198-410 ms. Considering the drawbacks mentioned above, this study aimed to determine a technical model of temporal regulation of movements and returner’s response, taking into account the continuous flow of information provided by the server. For this, a chronometric analysis of the returns of twelve players (six international and six national) of different sports level, that naturally responded by hitting their returns towards the targets, was performed. Actions of servers and returners were recorded with a Casio Exilim Pro Ex-F1 high speed camera (300 Hz) and then every 3.33 ms analysis was made frame by frame. Once the data of the videos were obtained, analyses were performed using one factor ANOVA test, ANCOVA with the speed of the serve as a covariate, U of Mann- Whitney and Pearson’s Chi-square test. As for the regulation of movement until the moment of serve, international players began their actions before national players, which could indicate that they were better prepared to execute movements reflecting the level of their expertise. International players began raising the rear foot at -293 ms and national players at -202 ms. All these actions were being linked to each other and it was at the moment of impact of the server when the receivers demonstrated a remarkable perceptual-motor coordination. Therefore, international players took off and started their flight just -6.5 ms before the serve and national players did the same somewhat later: +19.5 ms after the serve. Along the timeline, everything seems to indicate that the information used by returners interact with each other; early information which is less reliable to anticipate or move before, and later information more reliable appears to regulate the timing of actions. Returners of international and national levels anticipated at spatial level in a low percentage (7.7% vs. 13.6%) and in similar times (-127 vs. -118 ms) suggesting that the use of early and less reliable optical variables is only produced on rare occasions. Moreover, these data relate to a precise response as both international and national players showed a high percentage of success in responding (95.4% vs. 96.7%). It had been noted that international and national players would differ in the time the fall (i.e., landing) of the first foot of the split-step, however, this effect was not found (128 vs. 135 ms). No differences in the percentage of fall with the opposite foot to the direction of the ball (58% vs. 62%) were found. Where the two groups differed was in the time of the fall of the second foot (147 vs. 168 ms). This difference of 21 ms was crucial and it was a proof of mayor speed of international players; suggesting that this action could be related to the onset time of response. Although international players have proven to be faster in relation to their functional capabilities, both groups did not differ in all variables related to LVMr. They did not use those precious milliseconds earned at the time of the fall of the second foot to respond as soon, since the LVMr of the upper limb was the same for both groups (179 vs. 174 ms). It is as if they had all the time in the world to continue to adjust their actions until the return itself. Furthermore, these long times suggest that in the vast majority of the returns, key information that determined the response was detected (pick-up) in moments close to the hit of the server and in the first part of the ball flight. It was also found that in general the LVMr is influenced by the type of information used. Thus, when taking into account the trials during which there was spatial anticipation, reflected in LVMr of the whole body, the times decreased (152 vs. 136 ms). On the other hand, there were occasions (13%) where both international and national players responded late, thus receiving aces (208 vs. 195 ms). It is quite possible that in these cases the players have had trouble to pick-up information, responding out of temporary margins of action, which affected their performance. The same could have occurred when both groups of players corrected upper limb movement after impact (17% vs. 10%), which increased the time to respond and to redirect the return towards the right side (208 vs. 205 ms). Moreover, international players scored lower movement times than the national players (509 vs. 531 ms), which was reflected in a shorter total response time (683 vs. 703 ms). Finally, as far as the performance of return is concerned, international players scored above the national players values (1.3 vs. 0.9).

Regulation of meiotic chromatin loop size by chromosomal position.

At meiotic prophase, chromatin loops around a proteinaceous core, with the sizes of these loops varying between species. Comparison of the morphology of sequence-related inserts at different sites in transgenic mice demonstrates that loop size also varies with chromosomal geography. Similarly, chromatin loop lengths differ dramatically for interstitially and terminally located hamster telomeric sequences. Sequences, telomeric or otherwise, located at chromosome termini, closely associate with the meiotic proteinaceous core, forming shorter loops than identical interstitial sequences. Thus, we present evidence that different chromatin packaging mechanisms exist for interstitial versus terminal chromosomal regions, which act separately from those operating at the level of the DNA sequence. Chromosomal position plays the dominant role in chromatin packaging.

Grain size analysis of ODP Site 162-984

Past changes in the freshwater balance of the surface North Atlantic Ocean are thought to have influenced the rate of deep-water formation, and consequently climate (Broecker and Denton, 1989, doi:10.1016/0016-7037(89)90123-3; Manabe and Stouffer, 1996; doi:10.1038/378165a0). Although water-mass proxies are generally consistent with an impact of freshwater input on meridional overturning circulation (Boyle and Keigwin, 1987, doi:10.1038/330035a0), there has been little dynamic evidence to support this linkage. Here we present a 25,000 year record of variations in sediment grain size from south of Iceland, which indicates vigorous bottom-water currents during both the last glacial maximum and the Holocene period. Together with reconstructions of North Atlantic water-mass distribution, vigorous bottom currents suggest a shorter residence time of northern-source waters during the last glacial maximum, relative to the Holocene period. The most significant reductions in flow strength occur during periods that have been associated with freshening of the surface North Atlantic. The short-term deglacial oscillations in bottom current strength are closely coupled to changes in Greenland air temperature, with a minimum during the Younger Dryas cold reversal and a maximum at the time of rapid warming at the onset of the Holocene. Our results support a strong connection between ocean circulation and rapid climate change.

Temporal precision of interceptive action: differential effects of target size and speed

The duration of movements made to intercept moving targets decreases and movement speed increases when interception requires greater temporal precision. Changes in target size and target speed can have the same effect on required temporal precision, but the response to these changes differs: changes in target speed elicit larger changes in response speed. A possible explanation is that people attempt to strike the target in a central zone that does not vary much with variation in physical target size: the effective size of the target is relatively constant over changes in physical size. Three experiments are reported that test this idea. Participants performed two tasks: (1) strike a moving target with a bat moved perpendicular to the path of the target; (2) press on a force transducer when the target was in a location where it could be struck by the bat. Target speed was varied and target size held constant in experiment 1. Target speed and size were co-varied in experiment 2, keeping the required temporal precision constant. Target size was varied and target speed held constant in experiment 3 to give the same temporal precision as experiment 1. Duration of hitting movements decreased and maximum movement speed increased with increases in target speed and/or temporal precision requirements in all experiments. The effects were largest in experiment 1 and smallest in experiment 3. Analysis of a measure of effective target size (standard deviation of strike locations on the target) failed to support the hypothesis that performance differences could be explained in terms of effective size rather than actual physical size. In the pressing task, participants produced greater peak forces and shorter force pulses when the temporal precision required was greater, showing that the response to increasing temporal precision generalizes to different responses. It is concluded that target size and target speed have independent effects on performance.

Systematic changes in the duration and precision of interception in response to variation of amplitude and effector size

The results of two experiments are reported that examined how performance in a simple interceptive action (hitting a moving target) was influenced by the speed of the target, the size of the intercepting effector and the distance moved to make the interception. In Experiment 1, target speed and the width of the intercepting manipulandum (bat) were varied. The hypothesis that people make briefer movements, when the temporal accuracy and precision demands of the task are high, predicts that bat width and target speed will divisively interact in their effect on movement time (MT) and that shorter MTs will be associated with a smaller temporal variable error (VE). An alternative hypothesis that people initiate movement when the rate of expansion (ROE) of the target's image reaches a specific, fixed criterion value predicts that bat width will have no effect on MT. The results supported the first hypothesis: a statistically reliable interaction of the predicted form was obtained and the temporal VE was smaller for briefer movements. In Experiment 2, distance to move and target speed were varied. MT increased in direct proportion to distance and there was a divisive interaction between distance and speed; as in Experiment 1, temporal VE was smaller for briefer movements. The pattern of results could not be explained by the strategy of initiating movement at a fixed value of the ROE or at a fixed value of any other perceptual variable potentially available for initiating movement. It is argued that the results support pre-programming of MT with movement initiated when the target's time to arrival at the interception location reaches a criterion value that is matched to the pre-programmed MT. The data supported completely open-loop control when MT was less than between 200 and 240 ms with corrective sub-movements increasingly frequent for movements of longer duration.

A worldwide correlation for exponential bed particle size variation in subaerial aqueous flows

The particle size of the bed sediments in or on many natural streams, alluvial fans, laboratory flumes, irrigation canals and mine waste deltas varies exponentially with distance along the stream. A plot of the available worldwide exponential bed particle size diminution coefficient data against stream length is presented which shows that all the data lie within a single narrow band extending over virtually the whole range of stream lengths and bed sediment particle sizes found on Earth. This correlation applies to both natural and artificial flows with both sand and gravel beds, irrespective of either the solids concentration or whether normal or reverse sorting occurs. This strongly suggests that there are common mechanisms underlying the exponential diminution of bed particles in subaerial aqueous flows of all kinds. Thus existing models of sorting and abrasion applicable to some such flows may be applicable to others. A comparison of exponential laboratory abrasion and field diminution coefficients suggests that abrasion is unlikely to be significant in gravel and sand bed streams shorter than about 10 km to 100 km, and about 500 km, respectively. Copyright (C) 1999 John Wiley & Sons, Ltd.

Seawater carbonate chemistry, sample density and Strongylocentrotus purpuratus size, filtering and respiration rate during experiments, 2011

Anthropogenic CO2 emissions are acidifying the world's oceans. A growing body of evidence is showing that ocean acidification impacts growth and developmental rates of marine invertebrates. Here we test the impact of elevated seawater pCO2 (129 Pa, 1271 µatm) on early development, larval metabolic and feeding rates in a marine model organism, the sea urchin Strongylocentrotus purpuratus. Growth and development was assessed by measuring total body length, body rod length, postoral rod length and posterolateral rod length. Comparing these parameters between treatments suggests that larvae suffer from a developmental delay (by ca. 8%) rather than from the previously postulated reductions in size at comparable developmental stages. Further, we found maximum increases in respiration rates of + 100 % under elevated pCO2, while body length corrected feeding rates did not differ between larvae from both treatments. Calculating scope for growth illustrates that larvae raised under high pCO2 spent an average of 39 to 45% of the available energy for somatic growth, while control larvae could allocate between 78 and 80% of the available energy into growth processes. Our results highlight the importance of defining a standard frame of reference when comparing a given parameter between treatments, as observed differences can be easily due to comparison of different larval ages with their specific set of biological characters.

When is bigger better? The effects of group size on the evolution of helping behaviours.

Understanding the evolution of sociality in humans and other species requires understanding how selection on social behaviour varies with group size. However, the effects of group size are frequently obscured in the theoretical literature, which often makes assumptions that are at odds with empirical findings. In particular, mechanisms are suggested as supporting large-scale cooperation when they would in fact rapidly become ineffective with increasing group size. Here we review the literature on the evolution of helping behaviours (cooperation and altruism), and frame it using a simple synthetic model that allows us to delineate how the three main components of the selection pressure on helping must vary with increasing group size. The first component is the marginal benefit of helping to group members, which determines both direct fitness benefits to the actor and indirect fitness benefits to recipients. While this is often assumed to be independent of group size, marginal benefits are in practice likely to be maximal at intermediate group sizes for many types of collective action problems, and will eventually become very small in large groups due to the law of decreasing returns. The second component is the response of social partners on the past play of an actor, which underlies conditional behaviour under repeated social interactions. We argue that under realistic conditions on the transmission of information in a population, this response on past play decreases rapidly with increasing group size so that reciprocity alone (whether direct, indirect, or generalised) cannot sustain cooperation in very large groups. The final component is the relatedness between actor and recipient, which, according to the rules of inheritance, again decreases rapidly with increasing group size. These results explain why helping behaviours in very large social groups are limited to cases where the number of reproducing individuals is small, as in social insects, or where there are social institutions that can promote (possibly through sanctioning) large-scale cooperation, as in human societies. Finally, we discuss how individually devised institutions can foster the transition from small-scale to large-scale cooperative groups in human evolution.

First reproductive and productive results on “Ribatejano pig”

The present trail aimed to study the effect of crossbreeding between Alentejano (AL) and Bísaro (BI) swine breeds (“Ribatejano pig”) on some reproductive and productive traits. Nine AL gilts and sows and six BI gilts were crossed with BI and AL boars, respectively. Mating and farrowing dates, prolificacy and litter size at 28d were registered for all sows. The pregnancy length of AL sows was shorter (111±0.4d vs 113.7±0.5d; p=0.002) than the observed on BI females. The BI gilts presented higher prolificacy rate than AL on both total born (11.0±1.0 vs 6.7±0.8; P=0.004) and born alive piglets (10.0±1.0 vs 6.7±0.8; p=0.026). The mortality rate was similar in both genotypes (p=0.255) being on average 12%, therefore at 28d after farrowing the litter size remained higher in BI sows (8.5±0.8 vs 6.1±0.6; p= 0.032). A subset of each genotype (4 gilts) was supervised during farrowing and lactation (until 28d) and piglets were weighed at birth, 24h and 28d of live. Farrowing length was not significantly different (p=0.253) between genotypes, averaging 97±22 min. When compared to ALBI (AL x BI) piglets, the BIAL (BI x AL) piglets were heavier at birth (1402±46g vs 1209±36g; p=0.002). Colostrum intake of piglets per kg of birth weight on the first 24h of life was similar between genotypes (p=0.735) being 289±15g for ALBI and 281±19g for BIAL piglets. The growth rate of piglets from birth to 28d and piglet weight at 28d was not different between genotypes (p=0.161 and p=0.091) averaging 195±6g and 6761±181g, respectively. Litter weight at 28d tended (P=0.06) to be higher on ALBI litters (56.6±4.0kg) than BIAL litters (43.2±4.0kg). This results obtained within the frame of Treasure project* are, at our knowledge, the first data of these crossbred piglets and could be used in future as reference for further studies and also for farmers that may try these cross on a commercial basis.