MONOPHASIC AND DIPHASIC DIGESTIVE CYCLES IN VENERUPIS DECUSSATA AND CHLAMYS VARIA

Feeding and digestive cycles in Chlamys varia and Venerupisdecussata are identified and shown to relate to a tidal cycle.The processes of digestion within the diverticula of the twospecies are very different. The tubules of Venerupis are synchronizedand exhibit a monophasic cycle completed within a twelve-hourperiod, facilitating feeding at the next cycle of the tide.The digestive process of Chlamys requires twenty-four hoursand to accommodate feeding at each twelve-hour tidal cycle thetubules within the diverticula exhibit two different digestivephases simultaneously so that a diaphasic cycle is apparent.The pH of the mantle cavity and regions of the digestive tractand the formation and dissolution of the crystalline style areshown to be related to the tidal cycle. ^*Department of Brewing and Biological Sciences, Heriot-WattUniversity, Chambers Street, Edinburgh, EH1 1HX. (Received 22 March 1978;     

The effects of tidal currents on the rhythm of feeding and digestion in Pecten maximus L.

The orientation of individuals in two populations of Pecten maximus L., from the west coast of Ireland, shows that they have a marked preference to face directly into a tidal flow. In both localities examined there was a reversal of tides and the members of the populations were divided equally for flood and ebb tides. Twenty-four hour in situ studies of the animals were made and at all stages of the tide individuals facing either due east or west were observed. A cyclical feeding pattern imposed by the reversal of tidal flow is proposed.The pH of various parts of the digestive tract was investigated and showed a wide range of values. The most acid region was that of the stomach. The variations in pH of all of the regions of the gut examined throughout a 24-h period closely followed the pattern of the tidal cycle.Histological analysis of the stomach and digestive diverticula of representative samples of scallops taken at regular intervals over the 24-h periods clearly indicated a diphasic pattern of digestion within the tubules of the digestive diverticula. A close correlation between the phases of intracellular digestion, the pH variations in different regions of the gut, and the tidal cycle indicate distinct feeding cycles in Pecten. Those scallops facing into the ebb current show the same diphasic patterns as those individuals facing the flood current, but are 6 h out of phase. The cycle is considered as a duplication of a diphasic feeding pattern. The tubules themselves undergo a digestive process similar to that in Lasaea rubra (Montagu). The digestive cells phagocytose food material, begin intracellular digestion, and increase in size until they obscure the lumina. The dispersal of waste material and residual bodies is accomplished dramatically by a dehiscence of the tubule cells together with a loss of both digestive and crypt cells. New tubules are regenerated from the apices of those tubules breaking down. In any section of the diverticula tubules in two different conditions are found. The cycle of digestion takes 24 h and in order to facilitate feeding at each 12-h tidal cycle the tubules are equally divided into two phases with one 12 h behind the other.     

Glycosidases in the american oyster,Crassostreavirginica Gmelin,digestive tract

Activities for the glycosidases, cellobiase, maltase and chitobiase toward p-nitrophenyl-derived substrates, were found in the digestive diverticula of Crassostrea virginica (Gmelin). Chitobiase activity was also detected in the crystalline style, but specific activity was higher in the digestive diverticula. No activity was detected for either cellobiase or maltase in the style at pH values of 4.0, 4.25, 5.0 or 6.0 in 50 mM sodium acetate buffer or 7.0 in 100 mM phosphate (K₂HPO₄) buffer. Implications for digestive physiology are discussed.     

THE FUNCTIONAL MORPHOLOGY OF THE DIGESTIVE SYSTEM OF LYONSIA HYALINA CONRAD, 1831 (BIVALVIA: ANOMALODESMATA: PANDOROIDEA)

The anatomy and histology of the digestive tract of the suspensionfeeding bivalve Lyonsia hyalina were examined using microdissectionand conventional light microscopy. Lyonsia hyalina has a typeIV stomach which contains a major typhlosole that does not penetratethe left pouch, as in other members of the Pandoroidea. Theventral and posterior sorting areas of the stomach are sitesof vigorous ciliary activity. The gastric shield is locatedon the left and posterior stomach walls, underlain by tall basophiliccells with microvilli that project into the gastric shield.The style sac and midgut are combined, and contain the morphologicalcell types A-D seen in other bivalves. Many ciliated cells ofthe digestive tract appear to have high densities of apicalmitochondria. The ducts within the digestive diverticula arelined by epithelia containing a conspicuous brush border. Bothcrypt cells and digestive cells exist in the digestive tubules.The presence of numerous fragmentation spherules throughoutthe digestive diverticula indicates that intracellular digestionoccurs there. (Received 21 April 1992; accepted 1 October 1992)     

Regeneration of radiation-damaged digestive tissues in juvenile Pacific oysters (Crassostrea gigas)

Juvenile Pacific oysters, Crassostrea gigas, were irradiated with 16 and 40 krad and their tissues examined histologically. Degenerative syndromes and tissue regeneration processes were determined for the stomach, gut, collecting ducts, and digestive tubules. Following degeneration, tissue regeneration was observed in the digestive tissues of most oysters exposed to 16 krad and in a limited number exposed to 40 krad. Regeneration was first observed in the digestive tubules and subsequently in the stomach, gut, and collecting ducts. Cellular repopulation of the digestive tubules involved epithelialization with large, undifferentiated crypt cells which then differentiated into functional secretory and absorptive cells. Regeneration in the stomach, gut, and collecting ducts was initiated by proliferative islands of small basophilic cells. Mitotic division of those cells and their subsequent differentiation into functional epithelial cells resulted in the rapid restoration and apparent recovery of the affected tissues. The results of these studies indicate that radioresistance of juvenile C. gigas may in part be due to the remarkably efficient regenerative mechanisms involved in replacing injured or lost digestive tissues.     

洞庭湖区社鼠消化道长度和质量的季节变化

对洞庭湖区社鼠（Niviventer confucianus）自然种群四季的消化道各器官的长度和质量进行了测定。结果表明。消化道指标季节变化明显。总体消化道含内容鲜质量、净鲜质量与干质量均具有显著的季节变化。以冬、春季较高。夏、秋季较低。消化道各器官的变化与总消化道不尽相同。其中,以胃的变化相对比较稳定。仅长度的变化达显著水平,以冬季最短,夏、秋季较长。这是动物为适应繁殖季节能量需求而增加摄食量的反应,也与夏秋季食物丰富度有关。小肠、盲肠、大肠的长度和质量指标（内容鲜质量、净鲜质量和干质量）以冬、春季较高。夏、秋季较低。这是该鼠对冬、春季低温和食物匮乏的适应性反应。总体来说。洞庭湖区社鼠为适应夏、秋季繁殖盛期能量需求增加的主要对策是增加摄食量;而为适应冬、春季低温环境和食物数量和质量减少的主要对策是增大消化道。提高消化效率。     

Changes in the digestive gland of the loliginid squid Sepioteuthis lessoniana (Lesson 1830) associated with feeding

Changes associated with feeding in the histological and cytological structure of the digestive gland of the loliginid squid Sepioteuthis lessoniana were examined, along with the nature of both the intracellular and extracellular enzymes produced by the gland. The timing of the release of the extracellular enzymes during the digestive cycle was also determined using a quantitative experimental program. Like that of all coleoid cephalopods, the digestive gland was characterised by one type of cell with several functional stages. As is the case for other loliginid squids, however, the digestive cells did not contain the large enzyme-carrying boules that characterise the digestive glands of most cephalopods. Instead, smaller secretory granules were found in the digestive cells and these may be the enzyme carriers. The prominent rough endoplasmic reticulum, large mitochondria and active Golgi complexes present in the digestive cells are characteristic of cephalopods and indicate a high metabolic activity. Like that of other cephalopods, endocytotic absorption of nutrients and intracellular digestion occurs in the digestive gland of this squid. From quantitative and qualitative examinations of structural changes in the digestive gland of S. lessoniana after feeding, a schedule of its function during the course of digestion was proposed. This indicated that digestion was very rapid, being completed in as little as 4 h in S. lessoniana. Extracellular digestive enzymes were only released after the first hour following feeding, which implies that they are stored in the stomach between meals to increase digestive efficiency.     

新疆干旱环境中褐家鼠消化道长度和重量的季节变化及其适应意义

动物消化道的形态与食性、食物质量以及动物对能量的需求密切关系。本文对新疆干旱环境中生存的褐家鼠在自然环境中消化道长度及重量进行了测定,结果表明,家栖的褐家鼠只有总消化道、小肠和大肠的长度有季节差异,冬、春季高于夏、秋季。同时发现捕自阿拉山口的褐家鼠胃长度和干重有季节差异。     

Feeding rhythms of the green-lipped mussel, Perna viridis (Linnaeus, 1758) (Bivalvia: Mytilidae) during spring and neap tidal cycles

Food availability and feeding responses of the green mussel Perna viridis were investigated for two complete tidal cycles during both spring and neap tides. Temporal changes in total particulate matter (TPM), particulate inorganic matter (PIM) and particulate organic matter (POM), were smaller during neap than spring tides. Seston characteristics at different times of a tidal cycle were compared for both spring and neap tides. Only during spring tides were TPM and PIM significantly higher at high tides while POM remained relatively constant (P>0.05). The clearance rate of the mussels underwent temporal variations with tides, and was a negative power function of TPM and a positive linear function of f (organic content), during both spring and neap tides. f was the key factor influencing filtration rate, organic ingestion rate, absorption rate and absorption efficiency. All feeding rates increased linearly with increases in organic content. Pseudofaeces were produced only during spring but not neap tides. Feeding rates and absorption efficiency were highest at low and lowest at high tides (P<0.01). There was no significant temporal change in the wet weight and protein content of the crystalline style with the tidal regime. For the digestive gland, alpha-amylase activity was higher at spring than at neap tides, and higher during high tides in a tidal cycle. The digestive gland cellulase activity did not change significantly with the tides. For the crystalline style, both the activity of cellulase and alpha-amylase were not significantly different (P>0.05) between spring and neap tides. Tidal rhythms in feeding and digestion in this species were likely controlled by temporal variations in food availability in the seawater. By adjusting feeding rates and enzymatic activities, absorption in Perna viridis remained constant, irrespective of the changes in food availability.     

Epithelial cell renewal in the digestive gland and stomach of mussels: season, age and tidal regime related variations

The natural variability in cell proliferation activity in the epithelium of the digestive gland and stomach was investigated in mussels, Mytilus galloprovincialis (Lmk), of different age and tidal level at different seasons. After treating mussels with the thymidine analogue bromodeoxyuridine (BrdU) for 6 hours, BrdU immunohistochemistry was performed every 2 hours for the next 36. The relative proportion of BrdU positive cells was quantified as BrdU labelling (per thousand). Marked seasonal differences were recorded in BrdU labelling, with much higher proliferating activity in summer than in autumn and winter. Cell proliferation seemed not to be significantly dissimilar between mussels of different age (size). In contrast, the digestive gland epithelium of mussels from intertidal and subtidal populations differed not only in the levels but also in the pattern of variation of BrdU labelling, which in intertidal mussels appeared to be modulated by photoperiod and tide, unlike in subtidal mussels, in which variations followed a circatidal pattern.     

Seasonal and tidal variability of environmental carbon related physico-chemical variables and inorganic C acquisition in Gracilariopsis longissima and Enteromorpha intestinalis from Los Toruños salt marsh (Cádiz Bay, Spain)

The seasonal and tidal variability of inorganic C acquisition mechanisms, photosynthesis, internal composition and growth were studied in two co-occurring macroalgae in Los Toruños salt marsh (Cádiz Bay), Gracilariopsis longissima and Enteromorpha intestinalis. This variability was monitored together with physico-chemical variables affecting carbon availability, photosynthesis, and growth. The environmental variables, such as light, temperature, pH, salinity, oxygen, alkalinity, dissolved inorganic carbon (DIC) and CO₂, displayed not only an expected seasonal cycle but also a daily (tidal) variability, with abrupt and rapid changes influenced by biological activities, physical variables, tidal state and tidal timing. In contrast to environmental variables, photosynthesis, pigments and C:N composition were affected by seasonal changes but not by tidal regimes, as organisms integrated these short-term fluctuations in physico-chemical variables. Photosynthesis, pigments and internal N composition were maximal in autumn and minimal in summer for both species. Growth showed a seasonal trend, displaying a summer drop with negative values. This response can be the result of extreme values of environmental variables (temperature, light, pH, nutrients, and the shortage of DIC) in summer, in comparison with higher growth rates in September onwards. The use of inhibitors of carbon acquisition in situ at normal DIC concentrations (2.2. mM) revealed species-dependent differences. While the external carbonic anhydrase (CA) activity showed a constitutive character in G. longissima, it showed little effect in E. intestinalis, which relies on internal CA activity. The 4, 4′-diisothiocyanatostilbene-2,2′-disulfonate (DIDS)-sensitive bicarbonate transport in G. longissima was effective in winter. In contrast, DIDS stimulated photosynthesis in summer, and relieved AZ inhibition. This response could suggest a stimulation of a H⁺ extrusion mediated-CO₂ transport in periods of low CO₂ availability.     

The fine structure of the midgut of mite Myobia murismusculi

The pattern of digestion in females of Myobia murismusculi was studied with light and electron microscopy. The midgut consists of a stomach and two pairs of blind caeca. The stomach is connected dorsoposteriorly with the excretory organ, that leads externally to an anal opening via the cuticle-lined rectum. No differences were found between the stomach and its caeca. The midgut epithelial cells are of a single type. Their fine structure and gut contents greatly vary depending on different physiological conditions of the mite. Four stages of digestion can be shown with electron microscope. Pino- and phagocytose takes place in the same cells. At an active stage of digestion numerous pinocytic canals were observed in the midgut cells. At each stage of the digestive cycle groups of flat cells are present in the midgut epithelium. They do not take part in the intracellular digestion of food material. Cytoplasmic processes from the underlying cells of coxal glands project into the midgut cells through the orifices in the gut basal lamina.     

Effects of body-size and season on digestive organ size and the energy balance of cockles fed with a constant diet of phytoplankton

Seasonal variation in size-dependence of seawater clearance rate, absorption efficiency, oxygen consumption, gill area, length of the crystalline style and dry weight of digestive gland was analyzed in cockles Cerastoderma edule from the Mundaka Estuary, Spain. Experimental determinations were performed monthly (from July 1998 to November 1999) in cockles being fed with Tetraselmis suecica (organic content: 87.84 ± 1.95%) at a concentration of 3 mm³/l for 3 days. Analysis of covariance reveals no seasonal differences in both size-dependence of seawater clearance rate and oxygen consumption, which were found to scale to dry body weight with mass-exponents of 0.56 and 0.62, respectively. No significant correlation was found between absorption efficiency and body weight. Mass-exponents for gill area, dry weight of the digestive gland and length of the crystalline style remained constant among seasons showing values of 0.62, 0.34 and 0.82, respectively. Seasonal trends for every physiological determination were calculated for a standard size (200 mg) cockle: standardized clearance rates and oxygen consumptions followed a similar trend with minimum values in winter (≈0.5 l/h and ≈100 μl O₂/h, respectively) and maximum values during spring–summer (≈1.7 l/h and ≈250 μl O₂/h, respectively), whereas absorption efficiency and food throughput time showed both the opposite pattern with highest values corresponding to winter months (≈50–60% and ≈5–6 h, respectively), and lowest (≈30% and ≈3–4 h, respectively) to summer–autumn. Scope for growth exhibited minimum values in winter followed by a rapid increase along the winter–spring transition, maximum values being attained in spring (May) and summer (July). Exponential decline of seasonal values of absorption efficiency associated to rising ingestion rates of organic matter presented an asymptotic minimum at 0.35. Absorption efficiency was positively related to food throughput time, whereas the latter fell to a minimum of 3.548 h with increasing food intake. So, maintenance of throughput time—and consequently absorption efficiency—along with enhanced filtering activity provided cockles with higher absorption rates improving scopes for growth registers during spring and summer. These dynamics might be explained as the consequence of the seasonal digestive adjustments in cockles, which, in fact, were found to increase the size of the digestive organs during that period.     

Impact of fiddler crab foraging and tidal inundation on an intertidal sandflat: season-dependent effects in one tidal cycle

Intertidal sandflats inhabited by fiddler crabs are ideal systems in which to study the effects of physical and biological processes. This study addressed two questions: (1) Do fiddler crab feeding and tidal inundation have measurable effects on the sandflat over one tidal cycle? (2) Does the sandflat change over the course of a year? In field exclusion/inclusion experiments, fiddler crabs reduced sediment organic content by 40%, Chlorophyll a levels by 20% and meiofaunal density by 60% in one tidal cycle. Effects were most pronounced in the spring when organic content and meiofaunal densities were maximal. Effects of foraging were not erased by the tide and accumulated over time. The sandflat had highest levels of all variables in spring and minimal levels during summer and fall. Crabs graze the sandflat to minimum levels in the spring. Due to crab foraging, the flat is barren during the summer and fall, and recovers during the winter when crabs are minimally active.     

Fish assemblages across a vegetation gradient in a restoring tidal freshwater wetland: diets and potential for resource competition

Marsh habitats have been the focus of recent worldwide restoration efforts due to their degradation and destruction as a result of human development. We assessed fish resource use at a naturally restoring marsh (Liberty Island, California, USA) by comparing diet composition, stomach fullness, normalized stomach weight, and diet overlap across a vegetation gradient. Fish were collected using gill nets and fyke nets at six sites during spring 2010, summer 2011 and winter 2012. We analyzed 392 stomachs from the eleven most abundant species collected. Prey composition and biomass varied seasonally for all fish species, but there were no notable differences across sites or seasons for stomach fullness or normalized stomach weight for most fish species. Results from non-metric multidimensional scaling (NMDS) and two-way analysis of similarities (ANOSIM) indicated minimal diet overlap between species (R?=?0.633, p?=?0.001) and seasons (R?=?0.413, p?=?0.001). Seasonal habitat and resource use across the vegetation gradient was species-specific. Small but significant spatial diet differences were detected for inland silverside, striped bass and bigscale logperch. Delta smelt exhibited seasonal diet differences by shifting from chironomids and zooplankton during spring, to amphipods and zooplankton during winter. More generally, fish maintained stomach fullness across all sites and seasons while maintaining minimal dietary overlap. Our study emphasizes the importance of tidal marshes as feeding habitat for several fish species, including the endangered delta smelt.     

Thermal relations of large crocodiles,Crocodylus porosus,free-ranging in a naturalistic situation

We monitored behaviour and environmental and body temperatures (T_b) in summer and winter in 11 salt-water crocodiles (Crocodylus porosus), of body mass 32 to 1010kg, free-ranging in naturalistic captivity in northern Australia. We found pronounced daily cycles in air and water temperatures in both winter (16 to 33 degrees C and 20 to 31degrees C, respectively) and summer (21 to 45 degrees C and 24 to 36 degrees C, respectively). In winter, crocodiles exposed themselves to the sun during the day and stayed in the water at night. In summer, they remained in the water during the day and emerged onto land at night. Body temperature showed a daily cycle the amplitude of which decreased with increasing mass, from 3.5 degrees C (mass 32kg) to 1.0 degrees C (660kg) in summer, and from 3.5 degrees C (42kg) to 1.4 degrees C (1010kg) in winter. Underlying the daily cycles in T_b were intermediate (10 to 13 day, tidal?) and seasonal cycles. Overall, values of modal T_b ranged from 25.1 to 28.7 degrees C in winter and from 28.4 to 33.6 degrees C in summer, trending upwards with body size. This pattern of continuous oscillations in T_b, with no daily plateau, is conspicuously different from that seen in crocodilians of small sizes and from the pattern usually regarded as typical of reptiles in general.     

Reproductive biology of Philodryas patagoniensis (Snakes: Dipsadidae) in south Brazil: male reproductive cycle

The male reproductive cycle of Philodryas patagoniensis in south Brazil was described through morpho‐anatomical and histological analysis of individuals deposited in zoological collections. Spermatogenesis occurred during late autumn–winter (June–September) and spermiogenesis occurred in spring–summer (October–March). The volume of the testes was smaller (quiescent) in winter, while the tubular diameter and the epithelial height of the seminiferous tubule were larger in summer (January–March). The ductus deferens presented spermatozoa all over the year and had no seasonal variation in diameter. The length of the kidney was larger in winter–spring (July–December), although the tubular diameter and epithelium height of the sexual segment of the kidney (SSK) were larger only in winter (July–September). Total testicular regression was observed in late autumn (May), simultaneously with the peak in SSK. Therefore, at the individual level, males exhibit a discontinuous cyclical reproduction. Considering the population level, the reproductive cycle is seasonal semisynchronous, with most of the individuals showing a reproductive peak in spring–summer (October–March). Here, we present evidence to support the importance of the microscopic approach to reproductive cycle studies. Finally, we discuss the intrinsic and extrinsic factors influencing P. patagoniensis reproductive patterns.     

Observations on the feeding mechanism,structure of the gut,and digestive physiology of the european lobster Homarus gammarus (L.) (Decapoda: Nephropidae)

Investigations have been made on the feeding mechanism, structure of the gut, and digestive physiology of the European lobster Homarus gammarus (L.).Ciné-photography has shown that the mandibles do not possess a masticatory function, merely serving to grip food morsels during the tearing process effected by the pulling action of the third maxillipeds. The remaining maxillipeds, together with the maxillae, then direct food fragments to the mouth for ingestion.Ingestion is facilitated by mucoid secretions discharged from the oesophageal tegumental glands; the glands do not appear to produce any enzymes which directly contribute to the digestive processes.The hepatopancreas is the principal organ concerned with digestion. It possesses a complex tubular organization in which sequential cellular differentiation culminates in the discharge of enzymes from the B-cells for extracellular digestion in the cardiac stomach. The enzymes are synthesized within vacuoles contained in the B-cell precursors (F-cells) and are secreted in three bursts of activity at 0–15 min, 1–2 h, and 3.5–5 h after a meal. The initial secretory phase is holocrine. Extracellular digestion involves esterases, arylamidases, and lipases; endopeptidases have not been positively identified by histochemical means despite the fact that Homarus is a carnivore. There is an intracellular digestive phase, not previously described in decapod crustaceans, at the 7–9 h post-ingestive stage in the hepatopancreatic R-cells which is effected by arylamidases and lipases.Various phosphatase enzymes have been identified in the hepatopancreatic cells. Acid and alkaline phosphatases are apparently concerned with several stages in the digestive cycle, including enzyme synthesis and secretion, and the absorption of digestive products. Adenosine triphosphatase activity is primarily associated with granules located in the distal R-cell cytoplasm; the possible significance of these granules in the elimination of metabolic wastes is discussed. Acid phosphatases and esterases are present in the midgut epithelium. The possibility of a passive uptake of material from the midgut lumen is considered.Faecal material in the hindgut is bound by mucoid secretions derived from the tegumental glands of this alimentary region; the mucus may also assist in defaecation.A complete digestive cycle in Homarus occupies ≈ 12 h.Food reserves in the gut consist principally of fat deposits in the R-cells, but minute amounts of glycogen can also be detected.No evidence of calcium, copper or ferric iron deposition in any part of the alimentary tract was found.     

The Dimensions, Characteristics and Functions of the Crystalline Style of Molluscs

Recent findings concerning the crystalline style dimensions in different species of bivalves under normal or unfavorable life conditions, which lead to the partial or total dissolution of the crystalline style, were analyzed. The rate of the crystalline style total dissolution in media with different pH values and under different conditions was assessed. A variety of functions and the main physical and chemical characteristics of the crystalline styles, as well as the role and contents of digestive enzymes in them, are discussed.     

CYCLIC ACTIVITY AND EPITHELIAL RENEWAL IN THE DIGESTIVE GLAND TUBULES OF THE MARINE PROSOBRANCH MAORICRYPTA MONOXYLA (LESSON)

Three cell types are present in tubules of the digestive glandof the marine prosobranch Maoricrypta monoxyla (Lesson). Histochemistry,and feeding and starvation experiments established that themain type, the digestive cell, is involved in endocytotic uptakeof food material from the lumen. Digestion of this materialoccurs within vacuoles, and indigestible material (indicatedby the dye trypan blue) accumulates in basal residual bodiescontaining lipofuscin pigment. Another cell type, the cryptcell, appears to secrete a glycoprotein, probably enzymaticin function. The third cell type contains large vacuoles butit has not been Established whether the contents are secretoryor excretory. The tubules undergo a cycle of digestive activity not relatedto the tidal cycle as in some marine molluscs, but probablyan indirect result of the feeding regime. The cycle begins withimmature tubules in which some endocytosis occurs. These developinto absorbing tubules involved mainly in food uptake. In maturetubules intracellular digestion occurs. At a later stage thetubules fragment to produce spherules which may conserve usefulmaterial, and finally they, disintegrate completely. Eighty per cent of the dividing cells in the epithelium occurin crypts, which are therefore considered to be the main sitesof epithelial renewal. The processes by which tubules may bereformed after breakdown are discussed (Received 28 September 1978;