Demonstration of the cellular expression of genes encoding molt-inhibiting hormone and crustacean hyperglycemic hormone in the eyestalk of the shore crab Carcinus maenas

Based on the amino acid sequence of the molt-inhibiting hormone of Carcinus maenas, two degenerated oligonucleotide primers were synthesized and used in the polymerase chain reaction. By use of complementary DNA of a library constructed from medulla terminalis-X-organ RNA of C. maenas as template, the specific complementary DNA between the primers was amplified, cloned and sequenced. This strategy revealed a DNA sequence for which the deduced amino acid sequence is identical to the recently published C. maenas molt-inhibiting hormone sequence as determined by Edman degradation. Visualization of messenger RNAs encoding molt-inhibiting hormone and crustacean hyperglycemic hormone in different perikarya of the X-organ was obtained using digoxigenin-labelled complementary RNA probes. Combination of immunocytochemical staining using polyclonal antisera against the native C. maenas neuropeptides and in situ hybridization performed on alternating sections confirmed the specificity of the reaction. The results show that there is no co-localization of molt-inhibiting hormone and crustacean hyperglycemic hormone at the messenger RNA and the protein level.     

Occurrence of immunoreactive enkephalins in a neurohemal organ and other nervous structures in the eyestalk of the shore crab,Carcinus maenas L. (Crustacea,Decapoda)

Summary Light-microscopical observations with immunofluorescence and peroxidase staining procedures revealed leu-enkephalin-like immunoreactivity in axon profiles of the sinus gland (SG) and in single small neurons in the optic ganglia of the eyestalk of Carcinus maenas. Electron microscopy of the SG showed reactivity to be associated with neurosecretory granules 82±23 nm in diameter. High performance liquid chromatography of SG-extracts revealed radioimmunoreactive substances with the retention times of synthetic met- and leu-enkephalin and met-enkephalin-Arg⁶-Phe⁷, respectively.     

Fine structure of the neurohemal sinus gland of the shore crab,Carcinus maenas,and immuno-electron-microscopic identification of neurosecretory endings according to their neuropeptide contents

Summary The sinus gland of the shore crab, Carcinus maenas, is a compact assembly of interdigitating neurosecretory axon endings abutting upon the thin basal lamina of a central hemolymph lacuna. Four types of axon endings are distinguishable by the size distribution, shape, electron density and core structure of their neurosecretory granules. One additional type of axon ending is characterized by electron-lucent vacuoles and vesicles. The axon profiles are surrounded by astrocyte-like glial cells. Various fixations followed by epoxy- or Lowicrylembedding were compared in order to optimize the preservation of the fine structure of the granule types and the antigenicity of their peptide hormone contents. By use of specific rabbit antisera, the crustacean hyperglycemic, molt-inhibiting, pigment-dispersing, and red-pigment-concentrating hormones were assigned to the four distinct granule types which showed no overlap of immunostaining. Epi-polarization microscopy and ultrathin section analysis of immunogold-stained Lowicrylembedded specimens revealed that immunoreactivity to Leu-enkephalin and proctolin is co-localized with moltinhibiting hormone immunoreactivity in the same type of granule. The size and core structure of the immunocytochemically identified granule types vary little with the different pretreatments but, in some cases, to a statistically significant extent. The present results are compared with those from earlier studies of sinus glands in different crustaceans. The methods of granule identification used in this study supplement the classical approach in granule typing; they are easier to perform and more reliable for the analysis of release phenomena in identified secretory neurons supplying the neurohemal sinus gland.     

Corrective note about R* cells of the digestive gland of the shore crab Carcinus maenas

In a previous paper, we described and discussed the possible functions of calcospherite-rich cells (R^* cells) in the digestive gland of the shore crab, Carcinus maenas. We recently realised that electron micrographs in this publication presented neither typical R^* cells nor their calcium phosphate granules. Indeed, our pictures showed spermatophores (filled with typical spermatozoa) that had contamined hepatopancreatic cell suspensions. As the present study indicates, this contamination is difficult to detect by optical microscopy because unstained R^* cells closely resemble spermatophores. However, morphological differences between these cell types appear clearly when observed by electron microscopy. The present paper describes a comparative study of cell populations isolated from female digestive glands; it validates our previous results obtained with male hepatopancreas and suggests a low containation of those male cell fractions by spermatophores.     

Occurrence of crustacean cardioactive peptide (CCAP) in the nervous system of an insect,Locusta migratoria

Summary Using a radioimmunoassay developed for the determination of crustacean cardioactive peptide (CCAP), immunoreactive material was detected in extracts of locust nervous tissue. Serial dilutions of a brain extract gave a displacement curve parallel to the CCAP standard curve. One locust nervous system was calculated to contain approximately 1.4 pmol CCAP-like material.In order to investigate whether the immunoreactive substance was similar or identical to the crustacean neuropeptide, isolation and complete characterization was carried out using 800 locust nervous systems. The isolation procedure consisted of pre-purification of the crude extract on a Sep-Pak cartridge, affinity chromatography on a column which was prepared by coupling of anti-CCAP antibody to CNBr-activated Sepharose, and reversed phase high performance liquid chromatography (HPLC). In the HPLC-profile immunoreactivity was confined to a single peak which co-chromatographed with authentic CCAP. The peptide was carboxymethylated and analyzed in an automated gas-phase sequencer. Its amino acid sequence, is identical to that of CCAP fromCarcinus maenas.Synthetic CCAP was tested on the isolated locust hindgut in vitro. The peptide proved to be a potent enhancer of gut contractions, with a significant effect being observable at concentrations of 10^–10 M. It is concluded that in the locust CCAP may function as a myotropic peptide.     

Organization of a phyllobranchiate gill from the green shore crab Carcinus maenas (Crustacea,Decapoda)

Summary The phyllobranchiate gills of the green shore crab Carcinus maenas have been examined histologically and ultrastructurally. Each gill lamella is bounded by a chitinous cuticle. The apical surface of the branchial epithelium contacts this cuticle, and a basal lamina segregates the epithelium from an intralamellar hemocoel. In animals acclimated to normal sea water, five epithelial cell types can be identified in the lamellae of the posterior gills: chief cells, striated cells, pillar cells, nephrocytes, and glycocytes. Chief cells are the predominant cells in the branchial epithelium. They are squamous or low cuboidal and likely play a role in respiration. Striated cells, which are probably involved in ionoregulation, are also squamous or low cuboidal. Basal folds of the striated cells contain mitochondria and interdigitate with the bodies and processes of adjacent cells. Pillar cells span the hemocoel to link the proximal and distal sides of a lamella. Nephrocytes are large, spherical cells with voluminous vacuoles. They are rimmed by foot processes or pedicels and frequently associate with the pillar cells. Glycocytes are pleomorphic cells packed with glycogen granules and multigranular rosettes. The glycocytes often mingle with the nephrocytes. Inclusion of the nephrocytes and glycocytes as members of the branchial epithelium is justified by their participation in intercellular junctions and their position internal to the epithelial basal lamina.     

Localization of pigment-dispersing hormone (PDH) immunoreactivity in the central nervous system of Carcinus maenas and Orconectes limosus (Crustacea), with reference to FMRFamide immunoreactivity in O. limosus

Summary By use of antisera raised against synthetic pigment-dispersing hormone (PDH) of Uca pugilator and FMRFamide, the distribution of immunoreactive structures in the central nervous system (CNS) of Carcinus maenas and Orconectes limosus was studied by light microscopy. In both species, a total of 10–12 PDH-positive perikarya occur amongst the anterior medial, dorsal lateral and angular somata of the cerebral ganglion (CG). In C. maenas, one PDH-perikaryon was found in each commissural ganglion (COG) and several more in the thoracic ganglion. In O. limosus, only four immunopositive perikarya could be demonstrated in the ventral nerve cord, i.e., two somata in the anterior and two in the posterior region of the suboesophageal ganglion (SOG). PDH-immunoreactive tracts and fiber plexuses were present in all central ganglia of both species, and individual axons were observed in the connectives. FMRFamide-immunoreactivity was studied in O. limosus only. Neurons of different morphological types were found throughout the entire CNS, including numerous perikarya in the anterior medial, anterior olfactory, dorsal lateral and posterior cell groups of the CG. Four perikarya were found in the COG, six large and numerous smaller ones in the SOG, and up to eight cells in each of the thoracic and abdominal ganglia. In each ganglion, the perikarya form fiber plexuses. Axons from neurons belonging to the CG could be traced into the ventral nerve cord; nerve fibers arising from perikarya in the SOG appeared to project to the posterior ganglia. In none of the structures examined colocalization of PDH- and FMRF-amide-immunoreactivity was observed.Dedicated to Prof. K.-E. Wohlfarth-Bottermann on the occasion of his 65th birthday     

Sex-specific mediation of foraging in the shore crab, Carcinus maenas

Experiments were conducted to investigate the sex-specific differences to feeding responses of the shore crab Carcinus maenas throughout the year. Results demonstrate that female shore crabs exhibit stronger feeding responses than males throughout the year with a significantly reduced feeding response in males during the summer months' reproductive season. We also studied the possible function(s) of the moulting hormone, 20-hydroxyecdysone (Crustecdysone) that has been described as a potential female-produced sex pheromone to initiate male reproductive behaviour in a number of crustaceans. We recently presented evidence that for shore crabs this is not the case and now show that the steroid is instead functioning as a sex-specific feeding deterrent protecting the moulting 'soft' female crabs. Whilst male shore crabs were deterred from prey (Mytilus edulis) and synthetic feeding stimulants glycine and taurine when these feeding stimulants were spiked with crustecdysone, intermoult female crabs were significantly less affected and rarely deterred from feeding. This sex specificity of the moulting hormone, in combination with the female sex pheromone, which has no anti-feeding properties, ensures that male crabs mate with soft-shelled, moulted females rather than engage in cannibalism, such as found frequently in cases when soft-shelled females are exposed to intermoult females.     

Immunocytochemical localization of pigment-dispersing hormone (PDH) and its coexistence with FMRFamide-immunoreactive material in the eyestalks of the decapod crustaceans Carcinus maenas and Orconectes limosus

Summary By use of a new antiserum, raised against synthetic pigment-dispersing hormone (PDH) from Uca pugilator, immunoreactive structures were studied at the light-microscopic level in the eyestalk ganglia of Carcinus maenas and Orconectes limosus. PDH-reactivity was mainly found in two types of neurons that were located between the medulla interna (MI) and the medulla terminalis (MT) in both species. Several additional perikarya were located in the distal part of the MI in O. limosus. In C. maenas, two to three PDH-positive perikarya were found in the region of the X-organ (XO) in the MT. Processes from single and clustered cells could be traced into all medullae of the eyestalk. Axons from the immunoreactive perikarya running between MI and MT form a larger tract that traverses the MT. Fibers from this tract give rise to extensive arborizations and plexuses throughout the proximal MT. A plexus containing very fine fibers is located at the surface of the MT in a position distal to the XO-area of C. maenas only. The proximal plexus also receives PDH-positive fibers through the optic nerve. PDH-perikarya in the cerebral ganglion may also project into the more distal regions of the eyestalk. Distal projections of the perikarya between the MI and MT consist of several branches. Most of these are directed toward the MI and ME (medulla externa) wherein they form highly organized, layered plexuses. One branch was traced into the principal neurohemal organ, the sinus gland (SG). In the SG, the tract gives off arborizations and neurosecretory terminals. It then proceeds in a proximal direction out of the SG, adjacent to the MT. Its further course could not be elucidated. The lamina ganglionaris (LG) receives PDH-fibers from the ME and fine processes from small perikarya located in close association with the LG in the distal part of the first optic chiasma. The architecture of PDH-positive elements was similar in both C. maenas and O. limosus. The distribution of these structures suggests that PDH is not only a neurohormone but may, in addition, have a role as a neurotransmitter or modulator. Immunostaining of successive sections with an FMRF-amide antiserum revealed co-localization of FMRFamideand PDH-immunoreactivities in most, but not all PDH-containing perikarya and fibers. The axonal branch leading to the SG and the SG proper were devoid of FMRFamide immunoreactivity.     

Distribution of a novel cardioactive neuropeptide (CCAP) in the nervous system of the shore crab Carcinus maenas

A radioimmunoassay (RIA) for the recently discovered crustacean cardioactive peptide (CCAP) has been developed and used to determine contents of CCAP in different parts of the nervous system of the shore crab Carcinus maenas. Immunoreactive material was detected throughout the nervous system. In contrast to the main ganglia which contained low levels of approximately 1.4 pmol CCAP/mg protein (brain and thoracic ganglion), a high concentration was found in a neurohemal structure, the pericardial organs (PO) (868 pmol/mg protein). A predominantly neurohormonal role of CCAP thus suggested is further supported by in vitro release studies. Incubation of POs in high (K+) saline showed that CCAP is secretable in considerable amounts by a Ca++-dependent release mechanism.     

Crustacean cardioactive peptide in the nervous system of the locust,Locusta migratoria: an immunocytochemical study on the ventral nerve cord and peripheral innervation

Summary Crustacean cardioactive peptide-immunoreactive neurons occur in the entire central nervous system of Locusta migratoria. The present paper focuses on mapping studies in the ventral nerve cord and on peripheral projection sites. Two types of contralaterally projecting neurons occur in all neuromers from the subesophageal to the seventh abdominal ganglia. One type forms terminals at the surface of the thoracic nerves 6 and 1, the distal perisympathetic organs, the lateral heart nerves, and on ventral and dorsal diaphragm muscles. Two large neurons in the anterior part and several neurons of a different type in the posterior part of the terminal ganglion project into the last tergal nerves. In the abdominal neuromers 1–7, two types of ipsilaterally projecting neurons occur, one of which gives rise to neurosecretory terminals in the distal perisympathetic organs, in peripheral areas of the transverse, stigmata and lateral heart nerves. Four subesophageal neurons have putative terminals in the neurilemma of the nervus corporis allati II, and in the corpora allata and cardiaca. In addition, several immunoreactive putative interneurons and other neurons were mapped in the ventral nerve cord. A new in situ whole-mount technique was essential for elucidation of the peripheral pathways and targets of the identified neurons, which suggest a role of the peptide in the control of heartbeat, abdominal ventilatory and visceral muscle activity.Abbreviations AG abdominal ganglia - AM alary muscle - AMN alary muscle nerve - CA corpus allatum - CC corpus cardiacum - dPSO distal perisympathetic organ - LHN lateral heart nerve - LT CCAP-immunoreactive lateral tract - NCA nervus corporis allati - NCC nervus corporis cardiaci - NM neuromer - PMN paramedian nerve - PSO perisympathetic organ - SOG subesophageal ganglion - VDM ventral diaphragm muscles - VNC ventral nerve cord     

The hair-peg organs of the shore crab,Carcinus maenas (Crustacea,Decapoda): Ultrastructure and functional properties of sensilla sensitive to changes in seawater concentration

Summary The hair-peg organs of the shore crab, Carcinus maenas, are modified hair-sensilla. A small hair shaft (peg) is surrounded by a tuft of solid cuticular bristles (hairs). Each hair-peg organ is innervated by 6 sensory neurons, 2 of which have scolopidial (type-I) dendrites. The outer segments of all dendrites pass through a cuticular canal extending to the articulated hair base in which the 2 type-I dendrites terminate. The other 4 (type-II) dendrites reach the clavate tip of the hair shaft and have access to a terminal pore and a large sickle-shaped aperture. Three inner and 8–12 outer enveloping cells belong to a hair-peg organ. The innermost enveloping cell contains a scolopale, which has desmosomal connections to the ciliary rootlets of the type-I dendrites. An inner and an outer sensillum lymph space are present. The ultrastructural features of the dendrites and the cuticular apparatus indicate that the hair-peg organs are bimodal sensilla, comprising 2 mechano- and 4 chemosensitive sensory neurons. Extracellular recordings from the leg nerve indicate that the chemosensitive neurons of the hair-peg organs respond to changes in seawater concentration in the physiological range of Carcinus maenas.Supported by the Deutsche Forschungsgemeinschaft (SFB 45/A1; W. Gnatzy)     

The ultrastructure of the sinus gland of Carcinus maenas (Crustacea: Decapoda)

Summary The sinus gland of Carcinus maenas consists of the swollen axonal endings of the neurosecretory cells of the major ganglia and acts as a storage release centre for the membrane bound neurosecretory material. These neurosecretory granules fall into five different types based on size and electron density. Their contents are released by exocytosis of the primary granules or smaller units budded from the primary granules.I thank Professor E. Naylor for his constant advice and Professor E. W. Knight-Jones, Department of Zoology, University College, Swansea, for the provision of laboratory facilities. I am grateful to the Science Research Council for the financial support. Finally, I thank the Electron Microscope Unit, Southampton General Hospital, where the work was completed.     

Structure and possible functions of the calcospherite-rich cells (R* cells) in the digestive gland of the shore crab CArcinus maenas

Summary R^*-cells of the digestive gland of Carcinus maenas have been investigated functionally and morphologically. A comparison of the capacity of separated cell suspensions to synthesize glycogen gave support to the hypothesis that R and R^* cells belong to the same cell line. The unexpected observation of R^* cells in gastric juice suggests that their release could represent a mode of redistribution of carbohydrate stores when the feeding activity of the crab is lower. Under electron microscopy, the calcospherites of R^* cells appeared to be surrounded by multiple membranous layers, and displayed tubular and vesicular structures in their core. High glucose-6-phosphatase (G6Pase) activity in the subcellular fraction that is enriched in calcospherites suggests that these membranes are derived from the endoplasmic reticulum, via a process in which the enzyme plays a key role. We propose that this is the way by which the R cell differentiates into R^* cell.     

Patterns and projections of crustacean cardioactive-peptide-immunoreactice neurones of the terminal ganglion of crayfish

Three distinct clusters of crustacean cardioactive-peptide-immunoreactive neurones occur in the terminal abdominal ganglion of the crayfish species Orconectes limosus, Astacus leptodactylus, Astacus astacus and Procambarus clarkii, as revealed by immunocytochemistry of whole-mount preparations and sections. They exhibit similar topology and projection patterns in all four studied species. An anterior ventral lateral and a posterior lateral cluster contain one small, strongly stained perikaryon and two large, less intensely stained perikarya, each showing contralateral projections. A posterior medial lateral cluster of up to six cells also contains these two types of perikarya. Whereas the small type perikarya belong to putative interneurones, the large type perikarya give rise to extensive neurohaemal plexuses in perineural sheaths of the third roots of the fifth abdominal ganglia, the connectives, the dorsal telson nerves, the ganglion itself, its roots and arteriolar supply. Thin fibres from these plexuses reach newly discovered putative neurohaemal areas around the hindgut and anus via the intestinal and the anal nerves, and directly innervate the phasic telson musculature. A comparison with earlier investigations of motoneurones and segmentation indicates that these three cell groups containing putative neurosecretory neurones may be members of at least three neuromeres in this ganglion. Crustacean cardioactive peptide released from these neurones may participate in the neurohumoral and modulatory control of different neuronal and muscle targets, thereby exceeding its previously established hindgut and heart excitatory effects.Abbreviations AG abdominal ganglion - adpl arteria dorsalis pleica - Ala arreria lateralis abdominalis - Asub arteria subneuralis - CCAP crustacean cardioactive peptide - CNS central nervous system - IR immunoreactive - LG lateral giant axon - LTr lateral tract - MDT medial dorsal tract - MG medial giant axon - M Tr medial tract - mcan musculus compressor ani - mfltp museulus flexor telsonos posterior - nan nervus ani (AG6 N5) - nant nervus anterior (AG6 N1, N2) - nia nervus intestinal anterior - nin nervus intestinalis (AG6 N7) - nip nervus intestinalis posterior - nteld nervus telsonos dorsalis (AG6 N6) - nielv nervus telsonos ventralis (AG6 N4) - nur nervus uropedalis (AG6 N3) - nven nervus ventralis (AG5 N3) - PIR peri-intestinal ring - PTF posterior telson flexor - VLT ventral lateral tract - VMT ventral medial tract - VNC ventral nerve cord - VIF ventral telson flexor - AVLC, PLC, PMLC anterior ventral lateral, posterior lateral, posterior medial lateral CCAP-immunoreactive cell cluster - A6AVC, A7AVC anterior ventral commissures - A7DCI dorsal commissure I - A7PVC posterior ventral commissure - A7SCII sensory commissure II - A7VCII, A7VCIII ventral commissures II and III of the sixth (A6) and seventh (A7) abdominal neuromer     

Peptidergic modulation of cardiovascular dynamics in the Dungeness crab,Cancer magister

Decapod crustacean pericardial organs contain extensive neurohormonal reserves which can be released directly into the haemolymph to act as physiological modulators. The present paper concerns the in vivo effects of two pericardial peptides, proctolin and crustacean cardioactive peptide, on cardiovascular dynamics in the crab Cancer magister. Infusion of proctolin into the pericardial sinus caused a slight decrease in heart rate concurrent with a large increase in cardiac stroke volume. It decreased haemolymph flow anteriorly through the paired anterolateral arteries and increased flow posteriorly and ventrally through the posterior aorta and sternal artery, respectively. The threshold for responses occurred at circulating concentrations of 10^-9 mol·l^-1, and haemolymph flows remained elevated for up to 30 min after peptide infusion. The effects of crustacean cardioactive peptide were less dramatic. Heart rate was not affected but a significant increase in stroke volume was observed. Crustacean cardioactive peptide increased haemolymph flow through the anterolateral arteries and increased scaphognathite rate. The threshold for crustacean cardioactive peptide activity was higher than for proctolin (10^-7 mol·l^-1 and 10^-6 mol·l^-1) but the responses to crustacean cardioactive peptide were of longer duration. The effects of proctolin on regional haemeolymph distribution in Cancer magister closely resemble the cardiovascular responses of this species when exposed to hypoxic conditions. These peptides may be implicated as cardiovascular regulators during environmental perturbations.     

Structural organization of two fast,rhythmically active crustacean muscles

Summary The organization of the flagellum abductor muscle and of a scaphognathite levator muscle of the green crab, Carcinus maenas, has been compared quantitatively using light and electron microscopy. These muscles are rhythmically active at relatively high frequencies and for long durations. Fibers of both muscles are interconnected to form fascicles of 50 or more fibers within which there is cytoplasmic continuity. A single muscle is made up of 8–12 fascicles. Individual fibers consist of a peripheral rind of densely packed mitochondria, a thick region of glycogen granules, and myofibrils arranged into scattered central islands. Less than half the volume-density of these muscles is contractile material, the balance being largely mitochondria and glycogen. The fibers within a muscle are structurally similar. They have short sarcomeres (about 2 m), thin to thick filament ratios of about 3:1, and junctions between the sarcoplasmic reticulum and the transverse tubules at the M line. Sarcoplasmic reticulum occupies about 10% of the myofibrillar volume-density. A well developed sarcoplasmic reticulum appears to underlie the capacities of these two muscles for high frequency contraction; extensive mitochondria and glycogen stores should confer fatigue resistance under both aerobic and anaerobic conditions.     

Crustacean cardioactive peptide-immunoreactive neurons in the ventral nerve cord and the brain of the meal beetle Tenebrio molitor during postembryonic development

Summary By use of an antiserum against the crustacean cardioactive peptide (CCAP) several types of bilaterally symmetrical neurons have been mapped quantitatively in the ventral nerve cord and in the brain of the meal beetle, Tenebrio molitor. The general architecture of these neurons was reconstructed from peroxidase-antiperoxidase-labelled whole-mount preparations. From the subesophageal to the seventh abdominal ganglia two types of neurons show a repetitive organization of contralateral projection patterns in each neuromere. The first type has few branches in the central neuropil and a distinct peripheral projection. The second type is characterized by an elaborate central branching pattern, which includes ascending and descending processes. Some of its peripheral branches were found to supply peripheral neurohemal areas. In the protocerebrum, 10 CCAP-immunoreactive neurons occur with projections into the superior median protocerebrum and the tritocerebrum. Immunopositive neurons were mapped in larval and various pupal stages, as well as in the adult. All types of identified neurons were found to persist throughout metamorphosis maintaining their essential structural and topological characteristics. The CCAP-immunoreactive neurons of T. molitor are compared with those described for the locust. Putative structural homologies of subsets of neurons in both species are discussed.     

Larval abundance and recruitment of Carcinus maenas L. close to its southern geographic limit: a case of match and mismatch

In the Ria Formosa lagoon (S. Portugal), Carcinus maenas larvae have been observed only during the cold part of the year (October–May), when the water temperature stays below 23 °C. Larvae suffer high mortality during day time (about 80% during 6 h). Hence, they will only develop outside the lagoon, as proved by the observation of zoea I stages exclusively. Main recruitment starts in April, i.e. much later than the first larvae appear. Small crabs prefer Zostera noltii patches and colonize other sites when the carapace width exceeds about 5 mm. A reserve of some small crabs is found in Zostera noltii patches nearly all year round. It is postulated that one of the key factors for successful propagation of the species is a match between physiological reactions (here in particular temperature effect on reproductive cycle and larval release), and the ecological conditions for larval survival and recruitment, such as predator impact, food availability or currents.