Colour morph related performance in the meadow grasshopper Chorthippus parallelus (Orthoptera,Acrididae)

Abstract 1. Polymorphism has been described for a number of herbivorous insects, but little is known about whether differences in body colour cause fitness differences. In Chorthippus parallelus, three main colour morphs occur, namely brown, green, and dorsally striped. 2. The present study examined colour morph abundances and morph‐related differences in body size, oviposition rate, and offspring numbers in females of C. parallelus collected in 15 montane grasslands. The study also examined the effect of plant species richness, composition, community productivity, and solar radiation on colour morph frequency and fitness. 3. The relative frequencies of the three colour morphs was 31.7% (brown), 33.1% (green), and 35.2% (dorsally striped), but the morphs were not evenly distributed across the 15 sites. 4. There was no effect of the habitat variables on the distribution of the green and the striped morph in the study sites, however 80% of the variation in the abundances of the brown morph was explained by plant species richness and composition. 5. Grasshopper size was equal among the morphs. Brown females laid significantly more egg pods than the green and dorsally striped morphs. There were no significant differences in offspring numbers among the colour morphs. 6. Body colour in C. parallelus seems to be a fitness‐relevant trait, raising the question of the evolutionary maintenance of polymorphism.     

METABOLIC ADAPTATION IN THE POLYMORPHIC LAND SNAIL ARIANTA ARBUSTORUM

The oxygen consumption of brown and yellow morphs of autumnand winter Arianta arbustorum was determined after adaptationto several laboratory microclimatic conditions. The data onmetabolism of autumn and winter collected snails were treatedstatistically by using factorial analysis of variance for eachseason. There are highly significant differences in the metabolicrate of autumn and winter animals resulting from the influencesof microclimatic elements like adaptation temperature, relativehumidity, environmental temperature as well as the effect ofphenotypic shell colour. There are also significant differencesin the interactions of these factors which relate the animalsto natural ecological influences. The energy utilization averagedabout 25 percent less by the brown morph than the yellow oneat 20–30°C. Comparisons were made between oxygen consumptionof the animals at these two seasons. The winter snails are characterizedby a depressed respiration rate, and significant lowering ofrespiratory Q₁₀. The results are discussed in the light of physiologicalselection by different responses of metabolic rate to microclimaticfactors on the two morphs. This is important in the maintenanceof genetic diversity and polymorphism in A. arbustorum. ^*Current address: Department of Zoology, University of El-Azhar,Egypt. (Received 23 March 1983;     

Shell colour variation in Bullia digitalis, a sand-dwelling, intertidal whelk (Gastropoda: Prosobranchia)

Bullia digitalis is an intertidal whelk that lives on sandy beaches in South Africa. It is highly variable in shell colour, with individuals varying from white to dark brown. This paper describes shell colour variation of B. digitalis at seven sites, along a 230 km coastline east of the Cape Peninsula. Seven colour forms were found: striped, violet, banded violet, banded brown, orange, pale yellow and white. These forms are probably genetically determined morphs. The striped form is the most common at all sites, constituting 53–62% of each sample. The violet is the second most common morph. Its frequencies are remarkably stable at 15–17%. The striped form blends well into the sandy environment and may therefore be of considerable cryptic value in concealing B. digitalis from predators. The violet form is highly conspicuous. Its stable frequency throughout the study area may represent a genetic balance that is not relevant to any visual advantages of the violet colour.     

The adaptive significance of colour pattern polymorphism in the Australian scincid lizard Lampropholis delicata

Females of Lampropholis delicata are dimorphic for colour pattern, the difference between morphs being the presence or absence of a distinct white mid-lateral stripe. A less distinct striped morph occurs also in males. We evaluated alternative hypotheses for the maintenance of this polymorphism by examining temporal and spatial variation in morph frequency, testing for differential selection among morphs using data on body size and reproductive traits from preserved specimens, and experimentally manipulating colour pattern in free-ranging lizards of both sexes, to assess the influence of the lateral stripe on survival rates. We found that the relative frequency of striped individuals varied among populations and decreased from north to south in both sexes, coincident with an increasing incidence of regenerated tails. Morph frequencies did not change through time within a population. Striped gravid females appeared to survive better and produced larger clutches than did non-striped females. In our experimental study, the relationship between survival and colour morph differed between the two sexes; males painted with a white lateral stripe had lower survival than control (brown stripe) males, but survival did not differ between striped and control females. The different response in the two sexes may be due partly to differences in temperature and microhabitat selection. We propose that the white lateral stripe decreases susceptibility to predators in gravid females but increases risk of predation in males, especially in combination with low temperatures. The polymorphism might be maintained by: (1) opposing fitness consequences of the stripe in males and females; (2) sex-specific habitat selection; and (3) gene flow in combination with spatial variation in relative fitness of the two morphs.     

Temperature-related diversity of shell colour in the intertidal gastropod Batillaria

The intertidal snail Batillaria exhibits remarkable variationin the shell colour within and among populations. Field studywas conducted to determine the factors in maintaining observedshell colour polymorphism. Geographical variations in shellcolour polymorphisms in B. attramentaria were significantlycorrelated with the temperature of the locality of the population.Darker morphs were predominant in colder regions, whereas lightermorphs increase their proportion in warmer regions. A consistentassociation was also found in B. multiformis that co-existedwith B. attramentaria. Strong predatory pressure imposed bydigenean trematode parasites was observed in B. attramentaria.However, it is unlikely to affect the colour variations, becauseno correlation exists between colour morphs and trematode parasitism.Although visual selection may also contribute to colour variationin Batillaria, no evidence is found for the existence of visualpredators that affect colour patterns of these snails. The deficitof variation in cold regions is possibly due to selection againstbrighter morphs, because bright colours reflect heat. Althoughdark shells absorb sunlight and may therefore be exposed tothe risks of overheating and drying up in a hot habitat, thedarkest morph was frequently observed in the warmer regions,suggesting that physical selection on the colour morphs canbe relaxed in the warmer environment. Our results suggest thatclimatic selection is one of the significant factors maintainingshell colour polymorphism in these intertidal snails. (Received 13 December 2006; accepted 14 April 2007)     

Shell-Boring Mechanism of the Gastropod, Purpura (Thais) lapillus: a Physiological Demonstration of the Role of Carbonic Anhydrase in the Dissolution of CaCO3

Measurement of carbonic anhydrase (CA) activity in homogenatesof accessory boring organs of the muricid gastropod,Purpuralapillus, by Meldrum and Roughton's manometric method showedthat CA is always present in boring as well as in inactive ABOs,but in variable amounts. Tests by the same method on whole ABOsin an isotonic solution were negative, proving that CA remainsintracellularly. Experiments in vivo on inhibition and activation demonstratedclearly that CA is responsible for demineralization of the valvesof lamellibranchs by Purpura:(a) at low concentrations of Diamox,partial inhibition of the enzymeoccurred, that is, the numberof complete holes decreased or disappeared and etchings increased;and at higher concentrations full inhibition took place. Inhibitionis reversed when snails are replaced in normal sea water; (b)action of pure CO₂ or mixtures of CO₂ and O₂ accelerated boring:in the optimal mixture, three times more boreholes were producedby snails than in the controls, and in about half the time.Under these conditions the reaction catalyzed by CA goes tothe right with hydration of CO₂ and is accompanied by releaseof H⁺ ions; in the presence of an increased concentration ofCO₂, the reaction is intensified and results in an additionalrelease of H⁺ ions. Consequently, destruction of CaCO₃ by theABO of Purpura in sea water enriched with CO₂ is accelerated. To identify the nature of the exchanged ion, we tested the effectof NaCl and KC1 on the boring mechanism in vivo by increasingthe amount of these salts in the seawater of snails with prey.In this case augmentation of the boring activity was noticed.These results suggest that the boring activity is accompaniedby complex ionic exchanges.     

SHELL SCARRING IN CALLIOSTOMA ZIZYPHINUM (PROSOBRANCHIA: TROCHIDAE) FROM STRANGFORD LOUGH, NORTHERN IRELAND

Pink and white shells of Calliostoma zizyphinum show both undamagedand damaged scars. White shelled individuals predominate inStrangford Lough, Northern Ireland where Calliostoma densitiesand scarring levels are high. No white shells were found intertidally,outside the entrance channel to the Lough. Significant differencesoccur in the size of Calliostoma shells between sampling sitesbut not between different colour morphs at each site. Howeverin pooled samples, pink individuals are significantly largerthan white individuals. When sites and morphs are consideredseparately, pink Calliostoma density is negatively correlatedwith water movement. At sites where pink Calliostoma occur,the percentage of pink shelled individuals is negatively correlatedwith total Calliostoma density. Damaged and undamaged scarring values per unit area of shell,show highly significant differences between sites and damagedscars are significantly higher in white individuals. Undamagedscars are not correlated with any of the environmental parametersrecorded, but are positively correlated with damaged scars suggestinga common causative factor. The level of damaged scarring ispositively correlated with crab/total Calliostoma ratio at allsites and where each colour morph was considered separately.Multiple regression analyses reveal that crab/Calliostoma ratiosaccount for 42% of the between site variation in damaged scars.Significantly higher levels of damaged scars are found at siteswith high crab densities and significantly larger individualsare found at sites where crab densities are intermediate invalue. The largest and most highly scarred individuals occur at siteswith most coarse substrata where Calliostoma are present intheir lowest densities. The higher scarring levels and smallersize of white individuals reflect either higher mortality orreduced growth in white shelled Calliostoma. (Received 25 November 1991; accepted 5 November 1992)     

Feeding of the Cymatiid Gastropod, Argobuccinum argus, in Relation to the Structure of the Proboscis and Secretions of the Proboscis Gland

Several gastropod families belonging to the superfamily Tonnaceaare known to bore into the calcareous shells of echinoids andbivalves, but little work has been done on the family Cymatiidae.New observations show that the cymatiid, Argobuccinum argus,which feeds on the tubicolous polychaete, Gunnarea capensis,possesses large proboscis glands which secrete 0.4 to 0.5N H₂SO₄;further, the secretion dissolves greater amounts of pure CaCO₃than does O.47N H₂SO₄, alone. A toxin is also present, whichabolishes responses to light and touch in several invertebrates. The proboscis is atypically pleurembolic, having a permanentproboscis sheath. Retraction is due to the muscles ot the probosciswall, and the contracted proboscis is not inverted. Unlike othermesogastropods, the Tonnacea have three pairs ofglands associatedwith the foregut: proboscis glands, true salivary glands, anda small, partly paired gland of unknown function immediatelyposterior to the buccal mass. It is concluded that because of an abundance of readily availablefood in the form of Gunnarea, A. argus does not drillinto animalsprotected by a calcareous shell. The presence of a CaCO₃ dissolvingmechanism indicates that the Cymatiidae ingeneral may be ableto employ this method of feeding.     

Fluctuating selection by water level on gynoecium colour polymorphism in an aquatic plant

Background and Aims

It has been proposed that variation in pollinator preferences or a fluctuating environment can act to maintain flower colour polymorphism. These two hypotheses were tested in an aquatic monocot Butomus umbellatus (Butomaceae) with a pink or white gynoecium in the field population.

Methods

Pollinator visitation was compared in experimental arrays of equivalent flowering cymes from both colour morphs. Seed set was compared between inter- and intramorph pollination under different water levels to test the effect of fluctuating environment on seed fertility.

Key Results

Overall, the major pollinator groups did not discriminate between colour morphs. Compared with the white morph, seed production in the pink morph under intermorph, intramorph and open pollination treatments was significantly higher when the water level was low but not when it was high. Precipitation in July was correlated with yearly seed production in the pink morph but not in the white morph.

Conclusions

The results indicated that the two colour morphs differed in their tolerance to water level. Our study on this aquatic plant provides additional evidence to support the hypothesis that flower colour polymorphism can be preserved by environmental heterogeneity.     

Evidence for the Involvement of Mitochondrial Respiration in Calcification in a Marine Coccolithophorid, Emiliania huxleyi

In the marine coccolithophorid, Emiliania huxleyi, CaCO₃ productionunder illumination showed a lag phase for about 3 h and thenincreased greatly. During the lag phase the rate of CaCO₃ productionin the light was similar to that in the dark. The productionof CaCO₃ in the dark was inhibited by the addition of 170 µMCCCP, 1 mM KCN and 1 mM SHAM. These results suggest that a littleproduction of CaCO₃ is supported by energy from mitochondrialrespiration, but that large amount of CaCO₃ production requiresphotosynthesis. ¹Present address: SDS Biotech K.K., Tsukuba Technology Center,Midorigahara 2-1, Tsukuba, Ibaraki, 300-26 Japan     

Accumulation and Utilization of Dissolved Inorganic Carbon by a Marine Unicellular Coccolithophorid, Emiliania huxleyi

The mechanism for utilization of dissolved inorganic carbon(DIC) was investigated in the marine unicellular calcareousalga Emiliania huxleyi, grown with constant aeration. The apparentK_0.5 (DIC), the concentration of DIC which attains one-halfof the maximum velocity of apparent photosynthesis, for photosyntheticevolution of O₂, measured under saturating light, was 5.5 mM(55 µM for CO₂) at pH 8.0 and 25°C. The value of K_0.5was not affected by inhibitors of carbonic anhydrase (CA), andan electrometric assay of CA showed that the enzyme was notinvolved in photosynthesis in this alga. The rate of photosyntheticfixation of ¹⁴C-DIC into acid-stable products was about 20 timeshigher than that into CaCO₃, irrespective of the external concentrationof DIC. In short-term experiments, ¹⁴C-DIC was usually incorporatedinto the internal pool of DIC (IIC) to concentrations up to13 to 16 times higher than that of the external DIC. CO₂ addedexternally was utilized mainly for fixation of CO₂ and accumulationof IIC. By contrast, HCO^-₃ was utilized mainly for productionof CaCO₃ and accumulation of IIC. Incorporation of ¹⁴C intoIIC was partially suppressed by DCMU or in darkness but itstransfer to CaCO₃ was unaffected. These results suggest thataccumulation of IIC in this alga, even under ordinary circumstances,is only partially responsible for increasing the efficiencyof utilization of DIC by photosynthetic fixation but may bemost useful for the production of CaCO₃. (Hydroxyethylidene) bisphosphonic acid, an inhibitor of thegrowth of CaCO₃ crystals, completely suppressed production ofCaCO₃. The accumulation of IIC was also partially suppressed,but photosynthetic fixation of CO₂ was enhanced. In a pulse-chaseexperiment with ¹⁴CDIC, ¹⁴C incorporated into IIC and CaCO₃in darkness was transferred to acid-stable products of photosynthesisin the light. These results suggest that ¹⁴C-DIC in IIC andpre-formed CaCO₃ may be useful sources of carbon for fixationof CO₂. (Received July 2, 1993; Accepted January 10, 1994)     

Mechanisms of Electron Flow through the QB Site in Photosystem II. 3. Effects of the Presence of Membrane Structure on the Redox Reactions at the QB Site

Reduction kinetics of chloro- or methyl-substituted benzoquinones(BQs) in spinach PSII membrane fragments or n-heptyl-rß-D-thioglucoside-extractedPSII core complexes (HTG-PSII) was studied and compared to thatin cyanobacterial PSII core complexes [Satoh et al. (1995) PlantCell Physiol. 36: 597]. It was found that the BQs accept electronsat two sites (the Q_B and PQ sites) in the both spinach preparationsas in the cyanobacterial preparation. Maximum turnover rates(V_max) and binding affinities (K_m) of the two sites were estimated.Comparison of the values in PSII membrane fragments with thosein HTG-PSII showed that removal of the membrane structure orlight-harvesting chlorophyll a/b protein complexes from thePSII core complexes had little effect on the characteristicsof the Q_B site, indicating that the HTG-PSII have the intactQ_B site and are good materials to study the site. The K_m andV_max values were comparable to those in the cyanobacterial preparation. Low affinity (high K_m values) and high V_max values of methyl-substitutedBQs to the Q_B site and almost the same rate of intrinsic electronflow through the Q_B site in the both spinach preparations furthersupport the hypothesis that the plastoquinone (PQ) moleculeat the Q_B site is not replaced by another PQ molecule but, afterreduction, only its head group goes out of the Q_B pocket, andcomes back after transferring the electrons and protons to afree PQ molecule [Satoh et al. (1993) Z. Naturforsch. 48c: 174]. (Received May 2, 1996; Accepted July 26, 1996)     

Role of Coccoliths in the Utilization of Inorganic Carbon by a Marine Unicellular Coccolithophorid, Emiliania huxleyi: a Survey Using Intact Cells and Protoplasts

The utilization of inorganic carbon and role of the coccolithswere investigated in intact cells and protoplasts of a marineunicellular calcareous alga, Emiliania huxleyi. Protoplastswith high photosynthetic activity were obtained by artificialdecalcification with 50 mM MES-NaOH (pH5.5). (1) The kineticsof the photosynthetic evolution of O₂ at various concentrationsof externally added NaHCO₃ were the same for intact cells andprotoplasts, indicating that the kinetic properties with respectto dissolved inorganic carbon (DIC) were not affected by thepresence or absence of the coccoliths on the cell surface. Double-reciprocalplots and plots of the concentration of substrate divided byvelocity (s/v) against the concentration of substrate (s) werebiphasic in the case of both intact cells and protoplasts. TheCO₂-utilization reaction was, therefore, considered to involvetwo processes with different values of K_m and V_max. From thekinetic analyses, K_m and V_max [µmoles O₂ (ml PCV)^–1h^–1] were deduced to be 92 µM and 76.3 for a "low-K_m"reaction and 4.1 mM and 252 for a "high-K_m" reaction, respectively.(2) In short-term (40-min) experiments, time courses of thetotal uptake of ¹⁴C-DIC and the incorporation of ¹⁴C into acid-stableproducts of photosynthesis and the internal pool of DIC, determinedas acid-labile compounds, under CO₂-limiting conditions (80µM) were very similar for intact cells and protoplasts.However, incorporation of ¹⁴C into CaCO₃ apparently occurredmore slowly in protoplasts than in intact cells. (3) In longterm (24-h) experiments, patterns of incorporation of ¹⁴C werealmost same for intact cells and protoplasts, with the exceptionthat the amount of ¹⁴C incorporated into CaCO₃ was much smallerin the former than the latter. The production of Ca¹⁴CO₃ increasedduring the course of 10 h after a 4-h lag. However, after 10h the level of Ca¹⁴CCO₃ started to decrease. The decrease wasaccompanied by an increase in ¹⁴C in the products of photosynthesis,suggesting that CaCO₃ was reutilized for the photosyntheticfixation of CO₂ and, therefore, that the coccoliths functionas sites of storage of DIC. However, the internal level of DICremained at the same level even after the supply of externalDIC has been almost completely depleted. (Received July 25, 1995; Accepted December 11, 1995)     

FOOD-RELATED CONDITIONING AND NEURONAL CORRELATES IN THE FRESHWATER SNAIL LYMNAEA STAGNALIS

The feeding response of Lymnaea stagnalis shows robust appetitiveconditioning to a novel chemostim-ulus (amyl acetate), whichis retained for more than 4 days. In scrru-isolated centralnervous system (CNS)-hp preparations taken from conditionedsnails, application of amyl acetate to lip tissue led to theonset of fictive feeding in 5/17 individuals, and excitationof an identified cerebral-buccal feeding intemeuron, CV1_a. ina further 8. Control group snails showed no response to amylacetate. Attempts were made to aversively condition L. stagnalis usinga food stimulus (sucrose) as the conditioned stimulus and eitheran electric shock or mechanical stimulus as the unconditionedstimulus. No conditioned response to sucrose (i.e. withdrawal)was seen following training. However, trained snails exhibiteda reduction in responsiveness which was manifest as inhibitionof the feeding response to sucrose and an increase in the timetaken to emerge from the shell following handling. This reducedresponsiveness was seen at 1 hr but was lost by 24 hr aftertraining, and was not observed in control group snails. Semi-isolatedCNS-hp preparations from experimental snails recorded 1-3 hrafter training exhibited either inhibition of fictive feedingor no response following application of sucrose. Control groupsnails showed the normal excitatory response to sucrose includinginduction of feeding motor output Differences in food-related learning between Lymnaea and othergastropod species are discussed in relation to lifestyle andfeeding strategies (Received 14 January 1996; accepted 3 May 1996)     

The Influence of Parental Color Morph on Mate Choice in the Cichlid Fish Cichlasoma nigrofasciatum

Cichlasoma nigrofasciatum of two different color morphs (striped or white), each of which had been taken care of from spawning up to 7 weeks of age by foster parents of the other color morph, were put together in a “free choice situation” and tested for their preference to spawn with a mate of the striped or of the white morph. The number of pairs of the same color (striped × striped, white × white) and of different color (“mixed” pairs, striped × white) was recorded and compared with the results of control fish. The latter had been taken care of from spawning up to 7 weeks of age by foster parents of the same color morph and were also tested in a free choice situation. The influence of parental color morph on mate choice was analysed in the first mate choice of naive animals (breeding for the first time). The stability over time of the preferences observed in the naive animals was then studied in the 18 months following the first mate choice experiment. Our results show that if young Cichlasoma nigrofasciatum had been taken care of by foster parents of the other color morph, but had grown up with siblings of their own color morph, approximately 9 % of the matings involved partners which had a different color (mixed pairs). In contrast, young which had been taken care of by foster parents of their own color morph and had grown up with siblings of their own color were completely segregated according to color during breeding (striped pairs or white pairs) and never formed mixed pairs. The importance of the influence of parental and sibling color morph on the preference to mate with a particular morph and, consequently, on speciation in cichlids in general are discussed.     

Gene frequency changes in Cepaea snails on the Marlborough Downs over 25 years

In 1985 we resurveyed the sites on the Marlborough Downs in southern England at which Cain and Currey in 1960/61 sampled Cepaea snails and thence introduced the term 'area effects' to describe large areas of uniform morph frequency. Some sites no longer harboured Cepaea and at others the species composition had changed, with a general spread of Cepaea hortensis at the expense of Cepaea nemoralis. The majority, however, permitted comparison of morph frequencies between the two surveys. In C. nemoralis, we detected a significant overall decrease in the frequency of the brown morph and estimate selection as 5–9% per generation. There was no apparent change in frequencies of banded morphs. In C. hortensis we detected a significant overall increase in the frequency of unbanded shells (1–3% selection per generation) and an almost significant decrease in the frequency of fusions within the banded class. There was insufficient colour polymorphism in C. hortensis to allow analysis of colour morph frequencies. These changes—all in the direction of reduced absorption of solar energy—resemble others detected in both species at other localities in southern England. Possible explanations include large-scale climatic effects and changes in vegetation.     

Body size influences differently the detectabilities of colour morphs of cryptic prey

Body size and coloration may contribute to variation in performance and fitness among individuals; for example, by influencing vulnerability to predators. Yet, the combined effect of size and colour pattern on susceptibility to visual predators has received little attention, particularly in camouflaged prey. In the colour polymorphic pygmy grasshopper Tetrix subulata (Linnaeus, 1758), females are larger than males, although there is a size overlap between sexes. In the present study, we investigated how body size and colour morph influenced detection of these grasshoppers, and whether differences in protective value among morphs change with size. We conducted a computer‐based experiment and compared how human ‘predators’ detected images of large, intermediate or small grasshoppers belonging to black, grey or striped colour morphs when embedded in photographs of natural grasshopper habitats. We found that time to detection increased with decreasing size, that differences in time to detection of the black, grey and striped morphs depended differently on body size, and that no single morph provided superior or inferior protection in all three size classes. By comparing morph frequencies in samples of male and female grasshoppers from natural populations, we also examined whether the joint effects of size and colour morph on detection could explain evolutionary dynamics in the wild. Morph frequency differences between sexes were largely in accordance with expectations from the results of the detection experiment. The results of the present study demonstrate that body size and colour morph can interactively influence detection of camouflaged prey. This may contribute to the morph frequency differences between male and female pygmy grasshoppers in the wild. Such interactive effects may also influence the dynamics of colour polymorphisms, and contribute to the evolution of ontogenetic colour change and sexual dichromatism. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 113 , 112–122.     

Calcification in the Green Alga Halimeda: II. THE EXCHANGE OF CA2+ AND THE OCCURRENCE OF AGE GRADIENTS IN CALCIFICATION AND PHOTOSYNTHESIS

In Halimeda cylindracea and H. tuna segments, the concentrationof CaCO₃, MgCO₃, protein, and chlorophyll, as well as segmentvolume and wet and dry weight, increase with ‘age’i.e. from the apex of a branch downwards. Photosynthetic andcalcification rates decrease with age as does the degree oflight stimulation of calcification. Studies of the exchange of ⁴⁵Ca between the Halimeda thallusand the sea water under various conditions showed that mostof the Ca exchange is between the cell walls, the aragonitecrystals, and the intercellular space. The cell wall has twodistinguishable phases with half-times (t_0?5) of 200 and 35min while the CaCO₃ has a rapidly exchanging phase with a t_0?5of approximately 6 min. The t_0?5 of the exchange of Ca betweenthe intercellular space and the external medium is estimatedat about 6 min, on the basis of uptake studies. If the integrityof the barrier between the intercellular space and the externalsea water, created by the adpressed peripheral utricles is destroyedthe t_0?5 is smaller (<<3 min). These kinetic studies as well as comparative measurements ofcalcification rates by both isotopic and chemical methods showthat the ⁴⁵Ca method for measuring calcification rates overestimatesthe calcification rate, due to binding of ⁴⁵Ca in the cell wallsand retention of ⁴⁵Ca in the intercellular space. The ¹⁴C methodgives more accurate results and has the further advantage ofallowing simultaneous measurement of the photosynthetic andcalcification rate on the same segment.     

Estimating chlorophyll a abundance from the 'phytoplankton colour' recorded by the Continuous Plankton Recorder survey: validation with simultaneous fluorometry

The green colour (measured with reference to standard colourcharts) of sections of the Continuous Plankton Recorder (CPR)filtering silk was compared with estimates of chlorophyll aconcentration derived from a fluorometer mounted on the CPRduring seven tows in the North Sea between February and May1991. After the green colour was assessed, the abundance ofphytoplankton cells on the filtering silks was quantified bymicroscope analysis. Data were collected for 115 10-nautical-milesamples over a total of seven cruises. For these 115 samples,there was only a weak (F_1.113 = 3.8, P = 0.05, r² = 0.03) positiverelationship between the colour of the filtering silk and thechlorophyll a concentration. However, when this comparison wasrestricted to four tows (68 10-nautical-mile samples) wherethe recorded phytoplankton cell abundance on the silks was verylow, there was a highly significant (F_1.66 /,69.1, P < 0.001,r² = 0.51) positive relationship between the silk colour andthe chlorophyll a concentration. By measuring the relative colourintensity of CPR standard colour categories and quantifyingthe individual variation in the assessment of colour, a theoreticalmodel was developed which pedicted that if the silks were colouredin direct proportion to the chlorophyll a concentration in thewater, then the expert r² for the relationship between silkcolour and chlorophyll a concentration would be 0.62. The greencolour recorded by the CPR survey was therefore identified asa quantitative index of chlorophyll a concentration, but onlywhen numbers of phytoplankton cells on the CPR silks are nothigh.