Practical application of spatial ecosystem service models to aid decision support.

Ecosystem service (ES) spatial modelling is a key component of the integrated assessments designed to support policies and management practices aiming at environmental sustainability. ESTIMAP ("Ecosystem Service Mapping Tool") is a collection of spatially explicit models, originally developed to support policies at a European scale. We based our analysis on 10 case studies, and 3 ES models. Each case study applied at least one model at a local scale. We analyzed the applications with respect to: the adaptation process; the "precision differential" which we define as the variation generated in the model between the degree of spatial variation within the spatial distribution of ES and what the model captures; the stakeholders' opinions on the usefulness of models. We propose a protocol for adapting ESTIMAP to the local conditions. We present the precision differential as a means of assessing how the type of model and level of model adaptation generate variation among model outputs. We then present the opinion of stakeholders; that in general considered the approach useful for stimulating discussion and supporting communication. Major constraints identified were the lack of spatial data with sufficient level of detail, and the level of expertise needed to set up and compute the models.

Assessing Regional-Scale Impacts of Short Rotation Coppices on Ecosystem Services by Modeling Land-Use Decisions.

Meeting the world's growing energy demand through bioenergy production involves extensive land-use change which could have severe environmental and social impacts. Second generation bioenergy feedstocks offer a possible solution to this problem. They have the potential to reduce land-use conflicts between food and bioenergy production as they can be grown on low quality land not suitable for food production. However, a comprehensive impact assessment that considers multiple ecosystem services (ESS) and biodiversity is needed to identify the environmentally best feedstock option, as trade-offs are inherent. In this study, we simulate the spatial distribution of short rotation coppices (SRCs) in the landscape of the Mulde watershed in Central Germany by modeling profit-maximizing farmers under different economic and policy-driven scenarios using a spatially explicit economic simulation model. This allows to derive general insights and a mechanistic understanding of regional-scale impacts on multiple ESS in the absence of large-scale implementation. The modeled distribution of SRCs, required to meet the regional demand of combined heat and power (CHP) plants for solid biomass, had little or no effect on the provided ESS. In the policy-driven scenario, placing SRCs on low or high quality soils to provide ecological focus areas, as required within the Common Agricultural Policy in the EU, had little effect on ESS. Only a substantial increase in the SRC production area, beyond the regional demand of CHP plants, had a relevant effect, namely a negative impact on food production as well as a positive impact on biodiversity and regulating ESS. Beneficial impacts occurred for single ESS. However, the number of sites with balanced ESS supply hardly increased due to larger shares of SRCs in the landscape. Regression analyses showed that the occurrence of sites with balanced ESS supply was more strongly driven by biophysical factors than by the SRC share in the landscape. This indicates that SRCs negligibly affect trade-offs between individual ESS. Coupling spatially explicit economic simulation models with environmental and ESS assessment models can contribute to a comprehensive impact assessment of bioenergy feedstocks that have not yet been planted.

Comparing bioenergy production sites in the Southeastern US regarding ecosystem service supply and demand.

Biomass for bioenergy is debated for its potential synergies or tradeoffs with other provisioning and regulating ecosystem services (ESS). This biomass may originate from different production systems and may be purposefully grown or obtained from residues. Increased concerns globally about the sustainable production of biomass for bioenergy has resulted in numerous certification schemes focusing on best management practices, mostly operating at the plot/field scale. In this study, we compare the ESS of two watersheds in the southeastern US. We show the ESS tradeoffs and synergies of plantation forestry, i.e., pine poles, and agricultural production, i.e., wheat straw and corn stover, with the counterfactual natural or semi-natural forest in both watersheds. The plantation forestry showed less distinct tradeoffs than did corn and wheat production, i.e., for carbon storage, P and sediment retention, groundwater recharge, and biodiversity. Using indicators of landscape composition and configuration, we showed that landscape planning can affect the overall ESS supply and can partly determine if locally set environmental thresholds are being met. Indicators on landscape composition, configuration and naturalness explained more than 30% of the variation in ESS supply. Landscape elements such as largely connected forest patches or more complex agricultural patches, e.g., mosaics with shrub and grassland patches, may enhance ESS supply in both of the bioenergy production systems. If tradeoffs between biomass production and other ESS are not addressed by landscape planning, it may be reasonable to include rules in certification schemes that require, e.g., the connectivity of natural or semi-natural forest patches in plantation forestry or semi-natural landscape elements in agricultural production systems. Integrating indicators on landscape configuration and composition into certification schemes is particularly relevant considering that certification schemes are governance tools used to ensure comparable sustainability standards for biomass produced in countries with variable or absent legal frameworks for landscape planning.

Shared developmental programme strongly constrains beak shape diversity in songbirds.

The striking diversity of bird beak shapes is an outcome of natural selection, yet the relative importance of the limitations imposed by the process of beak development on generating such variation is unclear. Untangling these factors requires mapping developmental mechanisms over a phylogeny far exceeding model systems studied thus far. We address this issue with a comparative morphometric analysis of beak shape in a diverse group of songbirds. Here we show that the dynamics of the proliferative growth zone must follow restrictive rules to explain the observed variation, with beak diversity constrained to a three parameter family of shapes, parameterized by length, depth and the degree of shear. We experimentally verify these predictions by analysing cell proliferation in the developing embryonic beaks of the zebra finch. Our findings indicate that beak shape variability in many songbirds is strongly constrained by shared properties of the developmental programme controlling the growth zone.

Biocompatibility and osteoconduction of macroporous silk fibroin implants in cortical defects in sheep.

The goal of the presented study was to compare the biocompatibility and cellular responses to porous silk fibroin (SF) scaffolds produced in a water-based (UPW) or a solvent based process (HFIP) using two different SF sources. For that reason, four different SF scaffolds were implanted (n=6) into drill hole defects in the cancellous bone of the sheep tibia and humerus. The scaffolds were evaluated histologically for biocompatibility, cell-material interaction, and cellular ingrowth. New bone formation was observed macroscopically and histologically at 8 weeks after implantation. For semiquantitative evaluation, the investigated parameters were scored and statistically analyzed (factorial ANOVA). All implants showed good biocompatibility as evident by low infiltration of inflammatory cells and the absent encapsulation of the scaffolds in connective tissue. Multinuclear foreign body giant cells (MFGCs) and macrophages were present in all parts of the scaffold at the material surface and actively degrading the SF material. Cell ingrowth and vascularization were uniform across the scaffold. However, in HFIP scaffolds, local regions of void pores were present throughout the scaffold, probably due to the low pore interconnectivity in this scaffold type in contrast to UPW scaffolds. The amount of newly formed bone was very low in both scaffold types but was more abundant in the periphery than in the center of the scaffolds and for HFIP scaffolds mainly restricted to single pores.

A natural O-ring optimizes the dispersal of fungal spores.

The forcibly ejected spores of ascomycete fungi must penetrate several millimetres of nearly still air surrounding sporocarps to reach dispersive airflows, and escape is facilitated when a spore is launched with large velocity. To launch, the spores of thousands of species are ejected through an apical ring, a small elastic pore. The startling diversity of apical ring and spore shapes and dimensions make them favoured characters for both species descriptions and the subsequent inference of relationships among species. However, the physical constraints shaping this diversity and the adaptive benefits of specific morphologies are not understood. Here, we develop an elastohydrodynamic theory of the spore's ejection through the apical ring and demonstrate that to avoid enormous energy losses during spore ejection, the four principal morphological dimensions of spore and apical ring must cluster within a nonlinear one-dimensional subspace. We test this prediction using morphological data for 45 fungal species from two different classes and 18 families. Our sampling encompasses multiple loss and gain events and potentially independent origins of this spore ejection mechanism. Although the individual dimensions of the spore and apical ring are only weakly correlated with each other, they collapse into the predicted subspace with high accuracy. The launch velocity appears to be within 2 per cent of the optimum for over 90 per cent of all forcibly ejected species. Although the morphological diversity of apical rings and spores appears startlingly diverse, a simple principle can be used to organize it.

Closely related bird species demonstrate flexibility between beak morphology and underlying developmental programs.

The astonishing variation in the shape and size of bird beaks reflects a wide range of dietary specializations that played an important role in avian diversification. Among Darwin's finches, ground finches (Geospiza spp.) have beaks that represent scaling variations of the same shape, which are generated by alterations in the signaling pathways that regulate growth of the two skeletal components of the beak: the prenasal cartilage (pnc) and the premaxillary bone (pmx). Whether this developmental mechanism is responsible for variation within groups of other closely related bird species, however, has remained unknown. Here, we report that the Caribbean bullfinches (Loxigilla spp.), which are closely related to Darwin's finches, have independently evolved beaks of a novel shape, different from Geospiza, but also varying from each other only in scaling. However, despite sharing the same beak shape, the signaling pathways and tissues patterning Loxigilla beaks differ among the three species. In Loxigilla noctis, as in Geospiza, the pnc develops first, shaped by Bmp4 and CaM signaling, followed by the development of the pmx, regulated by TGFßIIr, ß-catenin, and Dkk3 signaling. In contrast, beak morphogenesis in Loxigilla violacea and Loxigilla portoricensis is generated almost exclusively by the pmx through a mechanism in which Ihh and Bmp4 synergize to promote expansion of bone tissue. Together, our results demonstrate high flexibility in the relationship between morphology and underlying developmental causes, where different developmental programs can generate identical shapes, and similar developmental programs can pattern different shapes.

First detection of thiotrophic symbiont phylotypes in the pelagic marine environment.

Marine oligochaete and nematode thiotrophic symbionts (MONTS) form a phylogenetic cluster within the Gammaproteobacteria. For the symbionts that live on the nematode surface, environmental transmission is likely. However, until now, no free-living relatives have been found. In this study, we detected MONTS cluster members in offshore surface seawater of both the Caribbean and the Mediterranean Sea by PCR amplification of their 16S rRNA genes. This is the first evidence of members of this cluster in the pelagic environment. These may either be free-living forms of the symbionts or closely related, nonsymbiotic strains. In either case, their existence sheds light on the evolution of beneficial symbioses between shallow water invertebrates and sulfur-oxidizing bacteria.

Sequence variability of the pattern recognition receptor Mermaid mediates specificity of marine nematode symbioses.

Selection of a specific microbial partner by the host is an all-important process. It guarantees the persistence of highly specific symbioses throughout host generations. The cuticle of the marine nematode Laxus oneistus is covered by a single phylotype of sulfur-oxidizing bacteria. They are embedded in a layer of host-secreted mucus containing the mannose-binding protein Mermaid. This Ca(2+)-dependent lectin mediates symbiont aggregation and attachment to the nematode. Here, we show that Stilbonema majum-a symbiotic nematode co-occurring with L. oneistus in shallow water sediment-is covered by bacteria phylogenetically distinct to those covering L. oneistus. Mermaid cDNA analysis revealed extensive protein sequence variability in both the nematode species. We expressed three recombinant Mermaid isoforms, which based on the structural predictions display the most different carbohydrate recognition domains (CRDs). We show that the three CRDs (DNT, DDA and GDA types) possess different affinities for L. oneistus and S. majum symbionts. In particular, the GDA type, exclusively expressed by S. majum, displays highest agglutination activity towards its symbionts and lowest towards its L. oneistus symbionts. Moreover, incubation of L. oneistus in the GDA type does not result in complete symbiont detachment, whereas incubation in the other types does. This indicates that the presence of particular Mermaid isoforms on the nematode surface has a role in the attachment of specific symbionts. This is the first report of the functional role of sequence variability in a microbe-associated molecular patterns receptor in a beneficial association.

Indirect land-use changes can overcome carbon savings from biofuels in Brazil.

The planned expansion of biofuel plantations in Brazil could potentially cause both direct and indirect land-use changes (e.g., biofuel plantations replace rangelands, which replace forests). In this study, we use a spatially explicit model to project land-use changes caused by that expansion in 2020, assuming that ethanol (biodiesel) production increases by 35 (4) x 10(9) liter in the 2003-2020 period. Our simulations show that direct land-use changes will have a small impact on carbon emissions because most biofuel plantations would replace rangeland areas. However, indirect land-use changes, especially those pushing the rangeland frontier into the Amazonian forests, could offset the carbon savings from biofuels. Sugarcane ethanol and soybean biodiesel each contribute to nearly half of the projected indirect deforestation of 121,970 km(2) by 2020, creating a carbon debt that would take about 250 years to be repaid using these biofuels instead of fossil fuels. We also tested different crops that could serve as feedstock to fulfill Brazil's biodiesel demand and found that oil palm would cause the least land-use changes and associated carbon debt. The modeled livestock density increases by 0.09 head per hectare. But a higher increase of 0.13 head per hectare in the average livestock density throughout the country could avoid the indirect land-use changes caused by biofuels (even with soybean as the biodiesel feedstock), while still fulfilling all food and bioenergy demands. We suggest that a closer collaboration or strengthened institutional link between the biofuel and cattle-ranching sectors in the coming years is crucial for effective carbon savings from biofuels in Brazil.

Molecular characterization of the symbionts associated with marine nematodes of the genus Robbea.

Marine nematodes that carry sulfur-oxidizing bacteria on their cuticle (Stilbonematinae, Desmodoridae) migrate between oxidized and reduced sand layers thereby supplying their symbionts with oxygen and sulfide. These symbionts, in turn, constitute the worms' major food source. Due to the accessibility, abundance and relative simplicity of this association, stilbonematids may be useful to understand symbiosis establishment. Nevertheless, only the symbiont of Laxus oneistus has been found to constitute one single phylotype within the Gammaproteobacteria. Here, we characterized the symbionts of three yet undescribed nematodes that were morphologically identified as members of the genus Robbea. They were collected at the island of Corsica, the Cayman Islands and the Belize Barrier Reef. The surface of these worms is covered by a single layer of morphologically undistinguishable bacteria. 18S rDNA-based phylogenetic analysis showed that all three species belong to the Stilbonematinae, although they do not form a distinct cluster within that subfamily. 16S rDNA-based analysis of the symbionts placed them interspersed in the cluster comprising the sulfur-oxidizing symbionts of L. oneistus and of marine gutless oligochaetes. Finally, the presence and phylogeny of the aprA gene indicated that the symbionts of all three nematodes can use reduced sulfur compounds as an energy source.

An Ultrasound Assisted Anchoring Technique (BoneWelding Technology) for Fixation of Implants to Bone - A Histological Pilot Study in Sheep.

The BoneWelding((R)) Technology offers new opportunities to anchor implants within bone. The technology melted the surface of biodegradable polymer pins by means of ultrasound energy to mould material into the structures of the predrilled bone. Temperature changes were measured at the sites of implantation in an in vitro experiment. In the in vivo part of the study two types of implants were implanted in the limb of sheep to investigate the biocompatibility of the method. One implant type was made of PL-DL-lactide (PLA), the second one was a titanium core partially covered with PLA. Healing period was 2 and 6 months, with 3 sheep per group. Bone samples were evaluated radiologically, histologically and histomorphometrically for bone remodeling and inflammatory reactions. Results demonstrated mild and short temperature increase during insertion. New bone formed at the implant without evidence of inflammatory reaction. The amount of adjacent bone was increased compared to normal cancellous bone. It was concluded that the BoneWelding((R)) Technology proved to be a biocompatible technology to anchor biodegradable as well as titanium-PLA implants in bone.

Comparison of computer assisted surgery with conventional technique for treatment of abaxial distal phalanx fractures in horses: an in vitro study.

OBJECTIVE: To (1) evaluate and compare computer-assisted surgery (CAS) with conventional screw insertion (conventional osteosynthesis [COS]) for treatment of equine abaxial distal phalanx fractures; (2) compare planned screw position with actual postoperative position; and (3) determine preferred screw insertion direction. STUDY DESIGN: Experimental study. SAMPLE POPULATION: Cadaveric equine limbs (n=32). METHODS: In 8 specimens each, a 4.5 mm cortex bone screw was inserted in lag fashion in dorsopalmar (plantar) direction using CAS or COS. In 2 other groups of 8, the screws were inserted in opposite direction. Precision of CAS was determined by comparison of planned and actual screw position. Preferred screw direction was also assessed for CAS and COS. RESULTS: In 4 of 6 direct comparisons, screw positioning was significantly better with CAS. Results of precision analysis for screw position were similar to studies published in human medicine. None of evaluated criteria identified a preferred direction for screw insertion. CONCLUSION: For abaxial fractures of the distal phalanx, superior precision in screw position is achieved with CAS technique compared with COS technique. CLINICAL RELEVANCE: Abaxial fractures of the distal phalanx lend themselves to computer-assisted implantation of 1 screw in a dorsopalmar (plantar) direction. Because of the complex anatomic relationships, and our results, we discourage use of COS technique for repair of this fracture type.

Computer-assisted surgery for screw insertion into the distal sesamoid bone in horses: an in vitro study.

OBJECTIVE: To compare the precision of computer-assisted surgery with a conventional technique (CV) using a special guiding device for screw insertion into the distal sesamoid bone in horses. STUDY DESIGN: In vitro experimental study. SAMPLE POPULATION: Cadaveric forelimb specimens. METHODS: Insertion of a 3.5 mm cortex screw in lag fashion along the longitudinal axis of intact (non-fractured) distal sesamoid bones was evaluated in 2 groups (8 limbs each): CV and computer-assisted surgery (CAS). For CV, the screw was inserted using a special guiding device and fluoroscopy, whereas for CAS, the screw was inserted using computer-assisted navigation. The accuracy of screw placement was verified by radiography, computed tomography, and specimen dissection. RESULTS: Surgical precision was better in CAS compared with CV. CONCLUSION: CAS improves the accuracy of lateromedial screw insertion, in lag fashion, into the distal sesamoid bone. CLINICAL RELEVANCE: The CAS technique should be considered for improved accuracy of screw insertion in fractures of the distal sesamoid bone.

An animal model in sheep for biocompatibility testing of biomaterials in cancellous bones.

BACKGROUND: The past years have seen the development of many synthetic bone replacements. To test their biocompatibility and ability for osseointegration, osseoinduction and -conduction requires their placement within bone preferably in an animal experiment of a higher species. METHODS: A suitable experimental animal model in sheep with drill holes of 8 mm diameter and 13 mm depth within the proximal and distal humerus and femur for testing biocompatibility issues is introduced. RESULTS: This present sheep model allows the placing of up to 8 different test materials within one animal and because of the standardization of the bone defect, routine evaluation by means of histomorphometry is easily conducted. This method was used successfully in 66 White Alpine Sheep. When the drill holes were correctly placed no complications such as spontaneous fractures were encountered. CONCLUSION: This experimental animal model serves an excellent basis for testing the biocompatibility of novel biomaterials to be used as bone replacement or new bone formation enhancing materials.

Clinical evaluation of the CRIF 4.5/5.5 system for long-bone fracture repair in cattle.

OBJECTIVE: To report clinical evaluation of the clamp rod internal fixator 4.5/5.5 (CRIF 4.5/5.5) in bovine long-bone fracture repair. STUDY DESIGN: Retrospective study. ANIMALS: Cattle (n=22) with long-bone fractures. METHODS: Records for cattle with long-bone fractures repaired between 1999 and 2004 with CRIF 4.5/5.5 were reviewed. Quality of fracture repair, fracture healing, and clinical outcome were investigated by means of clinical examination, medical records, radiographs, and telephone questionnaire. RESULTS: Successful long-term outcome was achieved in 18 cattle (82%); 4 were euthanatized 2-14 days postoperatively because of fracture breakdowns. Two cattle had movement of clamps on the rod. Moderate to severe callus formation was evident in 11 cattle 6 months postoperatively. CONCLUSIONS: Movement of clamps on the rod was recognized as implant failure unique to the CRIF. This occurred in cattle with poor fracture stability because of an extensive cortical defect. The CRIF system may not be ideal to treat metacarpal/metatarsal fractures because its voluminous size makes skin closure difficult, thereby increasing the risk of postoperative infections. CLINICAL RELEVANCE: CRIF cannot be recommended for repair of complicated long-bone fractures in cattle.

Assessment of stiffness and strength of 4 different implants available for equine fracture treatment: a study on a 20 degrees oblique long-bone fracture model using a bone substitute.

OBJECTIVE: To compare the mechanical properties of 4 stabilization methods for equine long-bone fractures: dynamic compression plate (DCP), limited contact-DCPlate (LC-DCP), locking compression plate (LCP), and the clamp-rod internal fixator (CRIF--formerly VetFix). STUDY DESIGN: In vitro mechanical study. SAMPLE POPULATION: Bone substitute material (24 tubes) was cut at 20 degrees to the long axis of the tube to simulate an oblique mid-shaft fracture. METHODS: Tubes were divided into 4 groups (n=6) and double plated in an orthogonal configuration, with 1 screw of 1 implant being inserted in lag fashion through the "fracture". Thus, the groups were: (1) 2 DCP implants (4.5, broad, 10 holes); (2) 2 LC-DCP implants (5.5, broad, 10 holes); (3) 2 LCP implants (4.5/5.0, broad, 10 holes) and 4 head locking screws/plate; and (4) 2 CRIF (4.5/5.0) and 10 clamps in alternating position left and right of the rod. All constructs were tested in 4-point bending with a quasi-static load until failure. The implant with the interfragmentary screw was always positioned on the tension side of the construct. Force, displacement, and angular displacement at the "fracture" line were determined. Construct stiffness under low and high loads, yield strength, ultimate strength, and maximum angular displacement were determined. RESULTS: None of the implants failed; the strength of the bone substitute was the limiting factor. At low loads, no differences in stiffness were found among groups, but LCP constructs were stiffer than other constructs under high loads (P=.004). Ultimate strength was lowest in the LCP group (P=.01), whereas yield strength was highest for LCP constructs (409 N m, P=.004). CRIF had the lowest yield strength (117 N m, P=.004); no differences in yield strength (250 N m) were found between DCP and LC-DCP constructs. Differences were found for maximum angular displacement at the "fracture" line, between groups: LPC

Comparison of computer-assisted surgery with conventional technique for the treatment of axial distal phalanx fractures in horses: an in vitro study.

OBJECTIVES: To compare the precision obtained with computer-assisted screw insertion for treatment of mid-sagittal articular fractures of the distal phalanx (P3) with results achieved with a conventional technique. STUDY DESIGN: In vitro experimental study. SAMPLE POPULATION: Thirty-two cadaveric equine limbs. METHODS: Four groups of 8 limbs were studied. Either 1 or 2 screws were inserted perpendicular to an imaginary axial fracture of P3 using computer-assisted surgery (CAS) or conventional technique. Screw insertion time, predetermined screw length, inserted screw length, fit of the screw, and errors in placement were recorded. RESULTS: CAS technique took 15-20 minutes longer but resulted in greater precision of screw length and placement compared with the conventional technique. CONCLUSION: Improved precision in screw insertion with CAS makes insertion of 2 screws possible for repair of mid-sagittal P3 fractures. CLINICAL RELEVANCE: CAS although expensive improves precision in screw insertion into P3 and consequently should yield improved clinical outcome.

Biocompatibility and resorption of a brushite calcium phosphate cement.

A hydraulic calcium phosphate cement with beta-tricalcium phosphate (TCP) granules embedded in a matrix of dicalcium phosphate dihydrate (DCPD) was implanted in experimentally created defects in sheep. One type of defect consisted of a drill hole in the medial femoral condyle. The other, partial metaphyseal defect was located in the proximal aspect of the tibia plateau and was stabilized using a 3.5 mm T-plate. The bone samples of 2 animals each per group were harvested after 2, 4, 6 and 8 weeks. Samples were evaluated for cement resorption and signs of immediate reaction, such as inflammation, caused by the cement setting in situ. Differences regarding these aspects were assessed for both types of defects using macroscopical, radiological, histological and histomorphometrical evaluations. In both defects the brushite matrix was resorbed faster than the beta-TCP granules. The resorption front was followed directly by a front of new bone formation, in which residual beta-TCP granules were embedded. Cement resorption occurred through (i) extracellular liquid dissolution with cement disintegration and particle formation, and (ii) phagocytosis of the cement particles through macrophages. Signs of inflammation or immunologic response leading to delayed new bone formation were not noticed at any time. Cement degradation and new bone formation occurred slightly faster in the femur defects.

Paraxial protocadherin coordinates cell polarity during convergent extension via Rho A and JNK.

Convergent extension movements occur ubiquitously in animal development. This special type of cell movement is controlled by the Wnt/planar cell polarity (PCP) pathway. Here we show that Xenopus paraxial protocadherin (XPAPC) functionally interacts with the Wnt/PCP pathway in the control of convergence and extension (CE) movements in Xenopus laevis. XPAPC functions as a signalling molecule that coordinates cell polarity of the involuting mesoderm in mediolateral orientation and thus selectively promotes convergence in CE movements. XPAPC signals through the small GTPases Rho A and Rac 1 and c-jun N-terminal kinase (JNK). Loss of XPAPC function blocks Rho A-mediated JNK activation. Despite common downstream components, XPAPC and Wnt/PCP signalling are not redundant, and the activity of both, XPAPC and PCP signalling, is required to coordinate CE movements.