Is the cask of facilitation ready for bottling? A symposium on the connectivity and future directions of positive plant interactions

The 2009 British Ecological Society''s Annual Symposium entitled ‘Facilitation in Plant Communities’ was held at the University of Aberdeen, Scotland, from 20 to 22 April 2009. This was the first ever international meeting dedicated to the rapidly expanding field of facilitation. The aim of the symposium was to assess the current ‘state-of-play’ by contrasting findings from different systems and by looking outwards in an attempt to integrate this field with other related fields. It was also aimed at understanding how knowledge of facilitation can help understand community dynamics and be applied to ecosystem restoration. The symposium identified several key areas where future work is likely to be most profitable.     

Positive interactions can increase size inequality in plant populations

1.  Large variation in the size of individuals is a ubiquitous feature of natural plant populations. While the role of competition in generating this variation has been studied extensively, the potential effects of positive interactions among plants, which are common in high-stress environments, have not been investigated.
2.  Using an individual-based 'zone-of-influence' model, we investigate the effects of competition, abiotic stress and facilitation on size inequality in plant monocultures. In the model, stress reduces the growth rate of plants, and facilitation ameliorates the effects of stress. Both facilitation and competition occur in overlapping zones of influence. We tested some of the model's predictions with a field experiment using the clonal grass Elymus nutans in an alpine meadow.
3.  Facilitation increased the size inequality of model populations when there was no density-dependent mortality. This effect decreased with density as competition overwhelmed facilitation. The lowest size inequality was found at intermediate densities both with the model and in the field.
4.  When density-dependent mortality was included in the model, stress delayed its onset and reduced its rate by reducing growth rates, so the number of survivors at any point in time was higher under harsh than under more benign conditions. Facilitation increased size inequality during self-thinning.
5.   Synthesis . Our results demonstrate that facilitation interacts with abiotic stress and competition to influence the degree of size inequality in plant populations. Facilitation increased size inequality at low to intermediate densities and during self-thinning.     

Role of "external facilitation" in implementation of research findings: a qualitative evaluation of facilitation experiences in the Veterans Health Administration

Background

Facilitation has been identified in the literature as a potentially key component of successful implementation. It has not, however, either been well-defined or well-studied. Significant questions remain about the operational definition of facilitation and about the relationship of facilitation to other interventions, especially to other change agent roles when used in multi-faceted implementation projects. Researchers who are part of the Quality Enhancement Research Initiative (QUERI) are actively exploring various approaches and processes, including facilitation, to enable implementation of best practices in the Veterans Health Administration health care system – the largest integrated healthcare system in the United States. This paper describes a systematic, retrospective evaluation of implementation-related facilitation experiences within QUERI, a quality improvement program developed by the US Department of Veterans Affairs.

Methods

A post-hoc evaluation was conducted through a series of semi-structured interviews to examine the concept of facilitation across several multi-site QUERI implementation studies. The interview process is based on a technique developed in the field of education, which systematically enhances learning through experience by stimulating recall and reflection regarding past complex activities. An iterative content analysis approach relative to a set of conceptually-based interview questions was used for data analysis.

Findings

Findings suggest that facilitation, within an implementation study initiated by a central change agency, is a deliberate and valued process of interactive problem solving and support that occurs in the context of a recognized need for improvement and a supportive interpersonal relationship. Facilitation was described primarily as a distinct role with a number of potentially crucial behaviors and activities. Data further suggest that external facilitators were likely to use or integrate other implementation interventions, while performing this problem-solving and supportive role.

Preliminary Conclusions

This evaluation provides evidence to suggest that facilitation could be considered a distinct implementation intervention, just as audit and feedback, educational outreach, or similar methods are considered to be discrete interventions. As such, facilitation should be well-defined and explicitly evaluated for its perceived usefulness within multi-intervention implementation projects. Additionally, researchers should better define the specific contribution of facilitation to the success of implementation in different types of projects, different types of sites, and with evidence and innovations of varying levels of strength and complexity.     

Facilitation by a dwarf shrub enhances plant diversity of human-valued species at high elevations in the Himalayas of Nepal

Facilitation is a global phenomenon that occurs when one species promotes the growth, survival, or reproduction of another species, mostly in stressful environments. However, the importance of facilitation by shrubs in maintaining plant community diversity is not well evaluated in the Himalayas, especially for the richness and conservation of medicinal and human-valued species. Therefore, we aimed to explore the facilitative role of a dwarf shrub species, Berberis angulosa, in maintaining plant composition and richness of human-valued species in the Langtang valley of Nepal's Himalayas. We censused plant species in open patches and beneath Berberis during monsoon and post-monsoon (dry) seasons at three elevations.Total species richness and richness of human-valued species were significantly higher inside the Berberis canopy than in gaps; the former being 39% and the latter 46% greater under shrubs than in open sites. Facilitation by Berberis shrubs promoted plant community diversity irrespective of season and elevation; however, higher differences in mean species richness for both total plant species and human-valued species during the dry season and at high elevation indicated increased facilitation intensity under more stressful conditions. The facilitative effect of Berberis shrubs increased, combining both seasons, overall plant diversity by 19% (total=105), and human-valued species by 16% (total=56). Our results show the importance of facilitation by nurse shrubs in structuring plant communities and protecting medicinal and socio-ecologically important plants, thus enriching ecosystem services in the Himalayas. These results suggest nurse plant species should be incorporated into conservation policies and management strategies for effective biodiversity conservation and sustainability, especially in the face of climate change.     

Facilitation at the sexually differentiated laryngeal synapse of Xenopus laevis

Under physiological conditions, the laryngeal synapse of male Xenopus laevis exhibits marked facilitation during repetitive nerve stimulation. The male laryngeal synapse is weak and requires facilitation to produce muscle action potentials and ultimately sound. The female laryngeal synapse is strong: muscle contractions are produced to single nerve stimuli. We sought to determine if laryngeal synapses of males and females also differ in their ability to facilitate. To measure facilitation, laryngeal muscle action potentials were suppressed either postsynaptically by bathing the preparation in saline containing curare or presynaptically by bathing the preparation in reduced calcium/elevated magnesium saline. Facilitation of postsynaptic potential amplitude or quantal content in response to paired pulses was measured in male and female larynges: there is no sex difference in paired pulse facilitation. Facilitation in response to trains of stimuli, in curare-blocked preparations, increased and reached plateau values more rapidly in females than in males, although the facilitation between the last and first pulses in the train was the same in the sexes. Thus, the sexually differentiated behavior of this synapse is controlled more by a sex difference in synaptic strength than by a sex difference in the ability to facilitate. Accepted: 14 June 1997     

Effects of mobile buffers on facilitation: experimental and computational studies

Facilitation is an important form of short-term plasticity that occurs in most synapses. At crayfish neuromuscular junctions, basal transmission and facilitation were significantly reduced after presynaptic introduction of "fast" high-affinity calcium buffers, and the decay of facilitation was accelerated. The existence of residual calcium during facilitation was also demonstrated. Computational modeling of three-dimensional buffered Ca(2+) diffusion and binding to secretory and facilitation targets suggest that the facilitation site is located away from a secretory trigger mediating exocytosis; otherwise, the facilitation site would be saturated by each action potential. Our simulations account for many characteristics of facilitation and effects of exogenous buffer, and suggest that facilitation is caused by residual calcium gaining access to a site distinct from the secretory trigger through restricted diffusion.     

Augmentation and facilitation of transmitter release. A quantitative description at the frog neuromuscular junction

Endplate potentials were recorded from frog and toad sartorius neuromuscular junctions under conditions of greatly reduced quantal contents. The magnitudes of augmentation increased with the duration and frequency of stimulation, often increasing at an accelerating rate during 10-20-s conditioning trains. The magnitudes of the first and second components of facilitation also increased, but reached apparent steady state values within the first few seconds of stimulation. These observations could be accounted for by assuming (a) that augmentation and the first and second components of facilitation arise from underlying factors in the nerve terminal that act to increase transmitter release; (b) that each nerve impulse adds an increment to each of the underlying factors; (c) that the magnitude of the increment typically increases during the train for augmentation but remains constant for the components of facilitation; and (d) that the underlying factors decay with first-order kinetics with time constants of approximately 7 s for augmentation and 60 and 500 ms for the first and second components of facilitation, respectively. The increments of facilitation added by each impulse were about twice as large in the toad as in the frog. Facilitation was described better by assuming a power relationship between the underlying factor and the observed facilitation than by assuming a linear relationship. Augmentation was described by assuming either a linear or power relationship.     

Neural Mechanism of Facilitation System during Physical Fatigue

An enhanced facilitation system caused by motivational input plays an important role in supporting performance during physical fatigue. We tried to clarify the neural mechanisms of the facilitation system during physical fatigue using magnetoencephalography (MEG) and a classical conditioning technique. Twelve right-handed volunteers participated in this study. Participants underwent MEG recording during the imagery of maximum grips of the right hand guided by metronome sounds for 10 min. Thereafter, fatigue-inducing maximum handgrip trials were performed for 10 min; the metronome sounds were started 5 min after the beginning of the handgrip trials. The metronome sounds were used as conditioned stimuli and maximum handgrip trials as unconditioned stimuli. The next day, they were randomly assigned to two groups in a single-blinded, two-crossover fashion to undergo two types of MEG recordings, that is, for the control and motivation sessions, during the imagery of maximum grips of the right hand guided by metronome sounds for 10 min. The alpha-band event-related desynchronizations (ERDs) of the motivation session relative to the control session within the time windows of 500 to 700 and 800 to 900 ms after the onset of handgrip cue sounds were identified in the sensorimotor areas. In addition, the alpha-band ERD within the time window of 400 to 500 ms was identified in the right dorsolateral prefrontal cortex (Brodmann''s area 46). The ERD level in the right dorsolateral prefrontal cortex was positively associated with that in the sensorimotor areas within the time window of 500 to 700 ms. These results suggest that the right dorsolateral prefrontal cortex is involved in the neural substrates of the facilitation system and activates the sensorimotor areas during physical fatigue.     

Presynaptic potentials and facilitation of transmitter release in the squid giant synapse

Presynaptic potentials were studied during facilitation of transmitter release in the squid giant synapse. Changes in action potentials were found to cause some, but not all, of the facilitation during twin-pulse stimulation. During trains of action potentials, there were no progressive changes in presynaptic action potentials which could account for the growth of facilitation. Facilitation could still be detected in terminals which had undergone conditioning depolarization or hyperpolarization. Facilitation could be produced by small action potentials in low [Ca++]o and by small depolarizations in the presence of tetrodotoxin. Although the production of facilitation varied somewhat with presynaptic depolarization, nevertheless, approximately equal amounts of facilitation could be produced by depolarizations which caused the release of very different amounts of transmitter.     

The importance of conspecific facilitation during recruitment and regeneration: A case study in degraded mangroves

Facilitation can affect positively the survival, growth and fitness of neighboring plants and is able to promote the establishment of plant species under harsh environmental conditions. This study is one of the few to focus on conspecific nurse plants enabling the establishment of tree followers and the generally unknown interplay between facilitation and competition during recruitment and establishment.Field studies were conducted in an hypersaline area in north Brazil where the hydrological regime was disturbed by road construction in 1974. Avicennia germinans shrubs were able to recolonize subareas.Three different stages of re-colonization were identified by satellite imagery based on the vegetation coverage and were defined as the early, middle, and late stages. Different plant parameters, including height and crown radius of individual trees, were measured, and all plants separating mangrove seedlings and larger plants were stem-mapped.The Hegyi index was adapted to measure local interaction intensity, considering both positive (facilitation) and negative (competition) interactions among neighboring plants. Spatial point pattern analyses were combined with the interaction index to obtain a better overview of the strength and importance of the plant interactions within the different recolonizing stages.The spatial patterns displayed aggregation for all plants in all developmental stages. This is supported by the interaction index calculated for seedlings. The index, however, provides an early signal for the switching of plant interactions from facilitation to competition as the developmental stage increased. This feature remains undetected in the spatial patterns because self-thinning processes are linked to individual growth processes and require more time to take place.This study broadens the ecological concept of nurse plants from hetero- to conspecific, including the importance and strength of plant interactions. The studied degraded areas are recolonizing naturally, and conclusions can be drawn for restoration projects, which usually do not take facilitation effects into account.     

The interplay between species' positive and negative interactions shapes the community biomass–species richness relationship

We used an individual-based spatially-explicit model to assess the role of facilitation and plant strategies in shaping the 'community biomass–species richness' relationship. Facilitation had few impacts on community's richness under both the most benign (high community biomass) and the most severe (low community biomass) environments where its intensity was weak. From medium to high environmental severity, facilitation increased community richness, because all plant strategies were facilitated. In contrast, from low to medium environmental severity facilitation decreased community richness, because only the most competitive species were facilitated, which induced a decrease in the richness of the stress-tolerant species overwhelming the increase in richness of the competitive species. Above all, our simulations show how 'strategy-dependent' interactions among species combine to shape the humped-back biomass–species richness relationship. It also demonstrates that facilitative effects might have long-term negative effects on species richness, which result is not included in current facilitation models.     

Facilitation allows plant coexistence in Cuban serpentine soils

Serpentine soils represent stressful habitats where plants have to cope with heavy metals, moisture limitation and low nutrient availability. We propose that facilitation is an important mechanism structuring plant communities under such stressful conditions. Facilitation has been shown to generate the spatial association of species, forming discrete vegetation patches of phylogenetically distant species. We measured these spatial and phylogenetic signatures left by facilitation in a serpentine plant community of central Cuba. Our results show that seedlings preferentially grow under plants of different species, and that adults are significantly aggregated into vegetation patches. In these patches, adults tend to co‐occur with distant relatives, ultimately forming phylogenetically diverse neighbourhoods. We discuss possible mechanisms explaining how species adapted to serpentine areas may be acting as nurses, reducing the stressful conditions for the establishment of other species.     

Previous Land Use Alters the Effect of Climate Change and Facilitation on Expanding Woodlands of Spanish Juniper

In Mediterranean–continental regions, changes in land use are leading to the expansion of valuable habitats like endemic Juniperus thurifera woodlands, but the impact of reduced rainfall, due to climate change, on this expansion remains uncertain. We assessed the early performance of J. thurifera in different global change scenarios with and without facilitation. Saplings were transplanted in three ecosystems types with different previous land use (woodlands, former agricultural fields, and former livestock pastures), microhabitats (open vs. understory of adult trees) and were subjected to two watering regimens. We characterized the abiotic environment and measured survival, growth and two ecophysiological parameters. Former livestock pastures were the least favorable ecosystem, where the nursing effect was greatest. Former agricultural fields had the highest survival, but were more sensitive than woodlands to water scarcity. Reduced rainfall decreased photochemical efficiency, particularly in the least favorable scenarios, but did not affect survival. Water use efficiency enhanced growth under the canopy, but not in the open, whereas photochemical efficiency enhanced growth and survival more in the open. Facilitation was critical for effective recruitment in the harshest scenarios: former livestock pastures and reduced rainfall. Comparison with previous studies suggests that establishment depends on infrequent wet episodes. In Mediterranean ecosystems, changes in land use and climate change are leading to woodland expansion due to the modulating effect of facilitation of the oncoming adverse drier conditions. Nevertheless, the positive effect of facilitation and the negative impact of aridity on tree recruitment are strongly influenced by previous land-use history.     

Plant Facilitation on a Mine Tailings Dump

Facilitation is a potentially useful tool in restoration efforts. We investigated the causes of facilitation in planted Picea mariana seedlings in an unvegetated mine tailings dump. Clusters of plants doubled the survival rate in the first growing season compared to single plants. In the first year mycorrhizal inoculation had no effect on survival, but by the second growing season only mycorrhizal inoculated plants survived, most of these being in plant clusters. This suggests that facilitation in this environment is partly the result of interactions with mycorrhizae.     

Facilitation of transmitter release at squid synapses

Facilitation is shown to decay as a compound exponential with two time constants (T1, T2) at both giant and non-giant synapses in squid stellate ganglia bathed in solutions having low extracellular calcium concentrations ([Ca++]o). Maximum values of facilitation (F1) were significantly larger, and T1 was significantly smaller in giant than non-giant synapses. Decreases in [Ca++]o or increases in [Mn++]o had variable effects on T1 and F1, whereas decreases in temperature increased T1 but had insignificant effects on F1. The growth of facilitation during short trains of equal interval stimuli was adequately predicted by the linear summation model developed by Mallart and Martin (1967. J. Physiol. (Lond.). 193:676--694) for frog neuromuscular junctions. This result suggests that the underlying mechanisms of facilitation are similar in squid and other synapses which release many transmitter quanta.     

Sheep blowfly strike risk and management in Great Britain: a survey of current practice

Abstract.  The methods used for the control of sheep blowfly strike (ovine cutaneous myiasis) and the farm management factors associated with strike prevalence were examined using data from questionnaire survey returns provided by 966 sheep farmers in Great Britain, based on the period between March 2003 and February 2004. Overall, 91% of participants treated prophylactically with insecticides against blowfly strike; 39% treated twice and 11% treated more than three times in the year. Insect growth regulators (IGRs) were the most commonly chosen product (40%), especially the IGR cyromazine. Only 12% of farmers opted to dip their sheep in organophosphate insecticide against fly strike and 2% of farmers reported applying inappropriate products against strike to their sheep, such as ivermectin or 'drenches'. Farmers worming their ewes more often were 0.8 times less likely to report blowfly strike, but those who wormed their lambs more often were 1.2 times more likely to report strike. Pure-breed flocks were 0.7 times less likely to record an outbreak of blowfly strike than cross-breed flocks. Strike was less likely in ewe flocks grazed at higher altitude; however, this relationship with altitude was not seen in lambs. The results show that insecticides remain the primary tool used by almost all farmers to prevent strike and that the type of insecticides used and means of application have altered dramatically over the past 15 years. However, the prevalence of strike has remained almost unchanged over this period. Clearly careful attention to the type and timing of insecticide application, in association with a detailed understanding of the husbandry factors that predispose sheep to higher strike risk, is essential to allow the optimal management of strike problems.     

Mechanisms of neurotransmitter release facilitation in strontium solutions

Mechanisms of neurotransmitter release facilitation were studied using electrophysiological recording of end-plate currents (EPC) and nerve ending (NE) responses after substitution of extracellular Ca ions with Sr ions at the frog neuromuscular junction. The solutions with 0.5 mM concentration of Ca ions (calcium solution) or 1 mM concentration of Sr ions (strontium solution) were used where baseline neurotransmitter release (at low-frequency stimulation) is equal. Decay of paired-pulse facilitation of EPC at calcium solutions with increase of interpulse interval from 5 to 500 ms was well described by three-exponential function consisting of early, first and second components. Facilitation at strontium solutions was significantly diminished due mainly to decrease of early and first components. At the same time, EPC facilitation with rhythmic stimulation (10 or 50 imp/s) at strontium solutions was significantly increased. Also more pronounced decrease of NE response 3rd phase, reflecting potassium currents was detected under rhythmic stimulation of 50 imp/s at strontium solutions comparing to calcium solutions. It was concluded that facilitation sites underlying first and early components had lower affinity to Sr ions than to Ca ions. The enhancement of frequency facilitation at strontium solutions is mediated by two mechanisms: more pronounced broadening of NE action potential and increase of bivalent cation influx due to feebly marked activation of Ca(2+)-dependent potassium current by Sr ions, and slower dynamics of Sr(2+) removal from NE axoplasm comparing to Ca(2+).     

Disentangling the effects of facilitation on restoration of the Atlantic Forest

Facilitation is an important ecological mechanism with potential applications to forest restoration. We hypothesized that different facilitation treatments, distance from the forest edge and time since initiation of the experiment would affect forest restoration on abandoned pastures. Seed and seedling abundance, species richness and composition were recorded monthly during two years under isolated trees, bird perches and in open pasture. Seed arrival and seedling establishment were measured at 10 m and 300 m from the forest edge. We sampled a total of 131,826 seeds from 115 species and 487 seedlings from 46 species. Isolated trees and bird perches increased re-establishment of forest species; however, species richness was higher under isolated trees. Overall, abundance and richness of seeds and seedlings differed between sampling years, but was unaffected by distance from the forest edge. On the other hand, species composition of seeds and seedlings differed among facilitation treatments, distance from the forest edge and between years. Seedling establishment success rate was larger in large-seeded species than medium- and small-seeded species. Our results suggest that isolated trees enhance forest re-establishment, while bird perches provide a complementary effort to restore tree abundance in abandoned pastures. However, the importance of seed arrival facilitation shifts toward establishment facilitation over time. Arriving species may vary depending on the distance from the forest edge and disperser attractors. Efforts to restore tropical forests on abandoned pastures should take into account a combination of both restoration strategies, effects of time and proximity to forest edge to maximize regeneration.     

Recurrent facilitation of spinal reflexes

Antidromic volleys in muscle nerves may facilitate monosynaptic reflexes originating from neighboring motoneurons. This facilitation has been studied in spinal cats. It is at its peak with a conditioning-test interval of 20 to 30 msec., and can last 50 to 100 msec. The threshold of facilitation is about the same as that of recurrent inhibition. Both phenomena appear to be activated by stimulation of the large motor axons. The latency of facilitation seems to be longer than that of recurrent inhibition by approximately 1 msec., suggesting the presence of at least one more synaptic delay. Facilitation often follows an inhibition of variable depth and duration. Frequently, however, the facilitation is not preceded by inhibition, and therefore it cannot be a rebound effect. The pharmacological properties of facilitation resemble those of recurrent inhibition. Dihydro-beta-erythroidine¹ partially blocks facilitation; the peak is decreased and occurs earlier, and the duration is shortened. Eserine increases the duration of facilitation and inhibition and sometimes enhances their magnitude. It is concluded that recurrent facilitation is mediated by the cholinergic axon collaterals, and that at least two interneurons are located between collateral and motoneuron. Possible mechanisms of facilitation are discussed.     

Recurrent Conditioning in the Cat Spinal Cord : Differential effect of meprobamate on recurrent facilitation and inhibition

The action of cumulative doses of meprobamate on antidromic conditioning has been studied in spinal cats. Recurrent facilitation is greatly reduced or completely abolished by total doses ranging from 210 to 400 mg./kg. The depth of recurrent inhibition is not affected in a consistent manner by meprobamate, but the duration of inhibition is markedly increased in all experiments. This differential action of meprobamate on facilitation and inhibition can be utilized to study conditioning effects consisting of combined inhibition and facilitation. If conditioning starts with an inhibitory phase, variable in duration, followed by facilitation, meprobamate depresses the facilitation and reveals an extended inhibitory curve. Facilitation, however, is not always accompanied by inhibition, since in some cases facilitation is depressed and no inhibition is uncovered. The results of these experiments are discussed in relation to the various types of conditioning that have been produced by antidromic stimulation.