Growth increments and biomineralization process in cephalopod statoliths

A study on morphological, structural and biochemical composition of Sepia officinalis and Loligo vulgaris statoliths and statocyst endolymph was undertaken with the aim of determining the major factors affecting the deposition process of statolith formation and to clarify the cause for the poor definition of the growth increments in S. officinalis statoliths. It is suggested that the different biochemical composition of the statocyst endolymph found in the two species accounts for distinct statolith crystallisation processes, which results in a different microstructure. This explains the better definition of growth increments in L. vulgaris statoliths comparing with those of S. officinalis. The protein content as well as Ca(2+) and Mg(2+) concentrations in the endolymph are more implicated in growth increments formation than Sr(2+) ion concentration. Moreover, the daily variations of the three factors mentioned, allowed us to formulate a working hypothesis to explain the daily deposition of growth increments: a dark ring (rich in organic matter) is deposited during daylight whereas a light ring (rich in CaCO(3)) during darkness. These results are discussed in the light of alternative hypotheses explaining the deposition mechanisms in statoliths.     

Age and growth of planktonic squids Cranchia scabra and Liocranchia reinhardti (Cephalopoda, Cranchiidae) in epipelagic waters of the central-east Atlantic

Statolith microstructure was studied in two abundant planktoniccranchiids, Cranchia scabra (56 specimens, 38–127 mm mantlelength, ML) and Liocranchia reinhardti (34 specimens, 99–205mm ML) sampled in epipelagic waters of the western part of theGulf of Guinea (tropical Atlantic). Growth increments were revealedin ground statoliths of both species. It was possible to distinguishtwo growth zones in statolith microstructure by their colourin reflected light of the microscope: the translucent postnuclearzone and pale white opaque zone. Assuming that growth incrementsin statoliths were produced daily, ages of the largest immatureC.scabra and L.reinhardti were 166 and 146 days, respectively.Both cranchiids are fast-growing squids with growth rates inlength resembling those of juveniles of tropical ommastrephidsand Thysanoteuthis rhombus. Liocranchia reinhardti grows faster:its growth rate in ML is approximately twice that of same-agedC.scabra. The life cycle of both cranchiids consists of twophases. During their epipelagic phase, C.scabra and L.reinhardtifeed and grow rapidly from paralarvae to immature young in theepipelagic waters, attaining 120–130 and 170–200mm ML by ages of 4–5 months, respectively. Then they changetheir life style to a deepwater phase.     

Statolith shape and microstructure as indicators of ontogenetic shifts in the squid Gonatus fabricii (Oegopsida, Gonatidae) from the Norwegian Sea

Statolith shape and microstructure were studied in 151 specimens of the common arctic squid Gonatus fabricii (7.3–322 mm pen length) collected in the southern Norwegian Sea. Statolith development and growth both comprised two main periods, which corresponded with the epipelagic and meso-bathypelagic ontogenetic periods of G. fabricii. During the epipelagic period (pen length range from 3 to 50–60 mm), statoliths quickly developed from the droplet-like form in early paralarvae to the pre-definite stage in juveniles (30–50 mm pen length). Paralarval and juvenile statoliths grew with high growth rates, and their microstructure contained narrow first-order growth increments. Three main growth zones (Z1, Z2, Z3) developed during this period, being well distinguished from each other by specific patterns of microstructure and separated from each other by distinct checks. During the meso-bathypelagic period (from 50–60 to 322 mm pen length), statoliths hardly changed their shape and grew very slowly. Only one growth zone (Z4) was formed within the statolith microstructure, characterized by disappearance of the first growth increments and formation of specific second-order bands. Each second-order band consisted of approximately seven first-order increments. If the assumption “one increment-one day” is true for G. fabricii, the squid would then be a slow-growing animal with a life span for both sexes not exceeding 2 years. Accepted: 25 May 1999     

Evaluation of the age of the red whelk Neptunea antiqua using statoliths, opercula and element ratios in the shell

Growth rings present in whole and sectioned statoliths were used to determine the age of red whelks Neptunea antiqua, from the North Sea. Validation of the periodicity of the rings was established in four whelks by comparing the number of statolith rings with the number of seasonal Mg:Ca ratio cycles present in shell calcium carbonate samples drilled sequentially from along the growth axis. There was exact correspondence between the number of growth rings and the number of element ratio cycles in two of the shells and a 1-year difference in the estimated age between the two methods in the other two shells, evidence which is strongly indicative of an annual periodicity of deposition to the statolith rings. The estimated age of the whelks using the statolith rings varied between 4 years (shell length 102 mm) and 17 years (shell length 148 mm). The age of the whelks ascertained from the statoliths was compared with age estimates from the number of adventitious layers in sectioned opercula. The number of adventitious layers in whelks from 51 to 148 mm shell length ranged between 1 and 12 years. No significant difference was observed between the number of strongly defined statolith rings and number of opercula adventitious layers.     

Statolith microstructure and age of early life stages of planktonic squids Galiteuthis phyllura and Belonella borealis (Oegopsida, Cranchiidae) from the northern North Pacific

Statolith morphology and microstructure were studied in twocommon species of panktonic cranchiid squids, Belonella borealis[four juveniles with mantle length (ML) 375–450 mm] andGaliteuthis phyllura (13 paralarvae and juveniles, ML 9–235mm),caught near the bottom and in pelagic layers over the continentalslope of Siberia in the northwest Bering Sea. The total numberof growth increments within the statoliths ranged from 277 to294 in B.borealis and from 10 to 209 in G.phyllura. Assumingthat these increments were produced daily, both species growrapidly in length (daily growth rate = 1.13mm day^–1 duringthe first 8–10 months of their juvenile phase in the mesopelagiclayers, prior to migration into deeper waters for maturation.     

Statolith shape and microstructure in studies of systematics, age and growth in planktonic paralarvae of gonatid squids (Cephalopoda, Oegopsida) from the western Bering Sea

The microstructure, morphology and ontogenetic development ofstatoliths and age and growth of 405 planktonic paralarvae and117 juveniles belonging to 10 species of gonatid squids (Cephalopoda,Oegopsida) were studied in the region of the continental slopein the western part of the Bering Sea (57°00'–61°30'N,163°00'E–179°20'W). The statolith microstructureof all species was characterized by the presence of a largedroplet-shaped nucleus and bipartite postnuclear zone dividedinto two by the first stress check, except for Berryteuthismagister which had only one stress check and an undivided postnuclearzone. In Gonatus spp., completion of development of the postnuclearzone coincided with full development of the central hook onthe tentacular club. Daily periodicity of statolith growth incrementswas validated by maintaining 13 paralarvae of the four mostabundant species in captivity. All species might be subdividedinto two groups based on statolith microstructure, i.e. specieswith a central position of the nucleus within the first statolithcheck (Gonatopsis spp., Egonatus tinro and B.magister) and specieswith the nucleus shifted to the inner side of the first statolithcheck (Gonatus spp.). Comparative analysis of statolith morphologyshowed that paralarval statoliths have species-specific charactersthat allowed the construction of keys to identify species ofgonatid paralarvae based on their statoliths. Analysis of paralarvalgrowth using statoliths revealed that these cold-water planktonicgonatid paralarvae have fast growth rates, attaining a mantlelength of 7–10 mm at 15–20 days and 20–25mm at 35–70 days.     

Structure of recent and fossil mysid statoliths (Crustacea,Mysidacea)

Statoliths of 61 Recent species representing all subfamilies of Mysidae were studied with special emphasis on internal structure. In addition 5 samples of fossil statoliths from Miocene deposits were examined. Species of Boreomysinae and Rhopalophthalminae show simple roughly spherical organic statoliths, with setae originating from the sensory cushion and anchored in the statolith with distal branches extending shortly below the surface. All other subfamilies possess mineralized statoliths of greater structural complexity, with differentiation in core and mantle, where each part may consist of up to three layers. Habitus is hemispherical to discoidal. External gross structures are dorsal tegmen, ventral fundus, and the ambitus forming the outer toroidal to semi-toroidal circumference. Setae penetrate the mantle through mineralic canals and insert on the surface of the core. As suggested by congeneric species of Schistomysis, there is no principal structural difference between statoliths mineralized with fluorite compared to vaterite. However, vaterite statoliths tend to be more often of moruloid appearance and are exceptional by showing a central conical hole (the hilum) or a central cavity in certain forms. These structures are typical of fossil calcite statoliths. In vaterite and fluorite statoliths, the mantle shows radially arranged (= spherulitic) crystal aggregates. Such arrangements are badly preserved in fossil calcite statoliths. In large extant statoliths, concentric structures, mainly in the form of superficial striation and/or concentric microstrata, are visible in coexistence with radial aggregates. Stratification is possibly due to stratified deposition of the nonmineralized gland product, while the spherulitic structure is indicative of subsequent radial growth of crystal aggregates. The structure of accessory fluorite statoliths in the statocyst of Mesopodopsis slabberi leads to the hypothesis that mantle material is formed by secretions of the caudal statocyst gland. After demineralization of fluorite, vaterite and calcite statoliths, an organic template remains showing most essential morphological features of the statolith. From this we conclude that the structure of the statolith is (almost) entirely matrix mediated. © 1993 Wiley-Liss, Inc.     

Micromanipulation of statoliths in gravity-sensing Chara rhizoids by optical tweezers

Infrared laser traps (optical tweezers) were used to micromanipulate statoliths in gravity-sensing rhizoids of the green alga Chara vulgaris Vail. We were able to hold and move statoliths with high accuracy and to observe directly the effects of statolith position on cell growth in horizontally positioned rhizoids. The first step in gravitropism, namely the physical action of gravity on statoliths, can be simulated by optical tweezers. The direct laser microirradiation of the rhizoid apex did not cause any visible damage to the cells. Through lateral positioning of statoliths a differential growth of the opposite flank of the cell wall could be induced, corresponding to bending growth in gravitropism. The acropetal displacement of the statolith complex into the extreme apex of the rhizoid caused a temporary decrease in cell growth rate. The rhizoids regained normal growth after remigration of the statoliths to their initial position 10–30 m basal to the rhizoid apex. During basipetal displacement of statoliths, cell growth continued and the statoliths remigrated towards the rhizoid tip after release from the optical trap. The resistance to statolith displacement increased towards the nucleus. The basipetal displacement of the whole complex of statoliths for a long distance (>100 m) caused an increase in cell diameter and a subsequent regaining of normal growth after the statoliths reappeared in the rhizoid apex. We conclude that the statolith displacement interferes with the mechanism of tip growth, i.e. with the transport of Golgi vesicles, either directly by mechanically blocking their flow and/or, indirectly, by disturbing the actomyosin system. In the presence of the actin inhibitor cytochalasin B the optical forces required for acropetal and basipetal displacement of statoliths were significantly reduced to a similar level. The lateral displacement of statoliths was not changed by cytochalasin B. The results indicate: (i) the viscous resistance to optical displacement of statoliths depends mainly on actin, (ii) the lateral displacement of statoliths is not impeded by actin filaments, (iii) the axially directed actin-mediated forces against optical displacement of statoliths (for a distance of 10 m) are stronger in the basipetal than in the acropetal direction, (iv) the forces acting on single statoliths by axially oriented actin filaments are estimated to be in the range of 11–110 pN for acropetal and of 18–180 pN for basipetal statolith displacements.Abbreviation CB cytochalasin B This work was supported by the Bundesminister für Forschung und Technologie, and by Fonds der Chemischen Industrie. We thank Professor Dr. A. Sievers (Botanisches Institut, Universität Bonn, Germany) for helpful discussions.     

A biomineralization study of the Indo-Pacific giant clam <Emphasis Type="Italic">Tridacna gigas</Emphasis>

The giant clam, Tridacna gigas, is an important faunal component of reef ecosystems of the Indo-Pacific region. In addition to its ecological role, shells of this bivalve species are useful bioarchives for past climate and environmental reconstructions. However, the biomineralization processes involved in shell aragonite deposition are insufficiently understood. Here, we present a study of the shell microstructure of modern specimens from Palm Island, Great Barrier Reef (GBR), Australia, and Huon Peninsula, Papua New Guinea (PNG), using a combination of petrography, scanning electron microscopy, electron backscatter diffraction, Raman spectroscopy and stable carbon isotope ratios. Daily growth increments were recognizable in all specimens through ontogeny, and counting these growth lines provides a robust specimen age estimate. For the internal layers, paired increments of organized aragonitic needles and compact, oblong crystals were recognized in a specimen from PNG, whereas specimens from GBR were composed of shield-like crystals that were not definable at the microscale. The combination of nutrient availability, rainfall and solar irradiance are likely to be the most significant factors controlling shell growth and may explain the observed differences in microstructure. The external layer, identical in all specimens, was composed of dendritic microstructure that is significantly enriched in ¹³C compared to the internal layer, suggesting different metabolic controls on layer deposition. We propose that the mineralization of the internal and external layers is independent from each other and associated with the activity of specific mantles. Future studies using T. gigas shells as bioarchives should consider the microstructure as it reflects the environment in which the individual lived and the differences in mineralization pathways of internal and external layers.     

AGE, GROWTH AND MATURATION OF THE SQUID ENOPLOTEUTHIS LEPTURA (OEGOPSIDA: ENOPLOTEUTHIDAE) FROM THE CENTRAL-EAST ATLANTIC

Fifty-nine specimens of the tropical epipelagic eno-ploteuthidEnoplotcuthis leptura were collected in the central-east Atlanticbetween 1986–1988. Statoliths were extracted from allspecimens (mantle length (ML) 4.1–92 mm) and processedunder the statolith ageing technique. The characteristic featureof statolith morphology in E. leptura is a sculpture of therostrum, which is covered by numerous tiny spines and knobs.In the ground statolith it was possible to distinguish fourmain growth zones consisting of narrow growth increments likethose in other squids studied. Allometric growth of statolithsversus ML is negative. E. leptura is a short-lived squid witha half-year life span. Growth rates of E. leptura are high atjuvenile stage (instantaneous rate of growth (G) of body weight(BW) 0.04–0.06). An early maturation of males (at age45–60 days) and females (at 80–90 days) causes asharp decrease of somatic growth of E. leptura, and mature squidhave low growth rates (G of BW - 0.OO3-O.0O5). Spawning takesplace between January and September with two peaks: in Januaryand in June-July. (Received 22 November 1992; accepted 15 February 1993)     

Unique proteins from the statoliths of Lolliguncula brevis (Cephalopoda: Loliginidae)

Information concerning the role of the organic matrix (OM) in statolith mineralization may contribute to resolving problems currently facing the use of increments in squid statoliths to estimate the age of individuals. A preliminary study aimed at purifying and characterizing the OM proteins from statoliths of the loliginid squid Lolliguncula brevis is described. Proteins extracted from the statoliths were separated into two fractions, insoluble and soluble in aqueous solutions. Gel electrophoresis indicated that the insoluble fraction is composed of at least eight major proteins, ranging in size from 25 to over 200 kDa. The five largest insoluble proteins were glycosylated, as shown by immunoassay. In contrast, the soluble fraction is composed of a single dominant protein of about 100 kDa, and two other major proteins of higher molecular weight. All three soluble proteins were glycosylated. Molecular weight and partial sequence data of peptides from five of the insoluble and one of the soluble proteins were used to search databases for possible homologs. No matches were found, suggesting that these proteins may belong to a class of hitherto undescribed OM compounds. Different proteins are involved in biomineralization processes in different organisms. We propose that the insoluble matrix proteins are responsible for regulating the appearance of increments in squid statoliths.     

阿根廷滑柔鱼两个群体间耳石和角质颚的形态差异

头足类硬组织具有稳定的形态特征、良好的信息储存以及抗腐蚀性等特点。根据2007年2—5月和2010年1—3月我国鱿钓船采集的阿根廷滑柔鱼样品,提取出625对耳石和787对角质颚,测量耳石10个形态数据和角质颚12个形态数据,通过除以胴长(Mantle Length,ML)校正后,对南巴塔哥尼亚群体(South Patagonic Stock,SPS)和布宜诺斯艾利斯-巴塔哥尼亚群体(Bonaerensis-Northpatagonic Stock,BNS)形态差异进行分析,并建立不同群体的判别函数。结果表明,BNS群体耳石和角质颚外形参数雌性大于雄性,而SPS群体则为雄性大于雌性。均数差异性检验认为,同一群体不同性别的耳石总长(Total StatolithLength,TSL)、最大宽度(Maximum Width,MW)、侧区长(Lateral Dome Length,LDL)、翼区长(Wing Length,WL)、翼区宽(WingWidth,WW)存在显著差异(P<0.05),同性别不同群体的MW、背侧区长(Dorsal Lateral Length,DLL)、吻侧区长(RostrumLateral Length,RLL)和WW存在显著差异(P<0.05)。而同一群体不同性别间角质颚的上头盖长(Upper Hood length,UHL)、上脊突长(Upper Crest length,UCL)、上喙长(Upper Rostrum length,URL)、上喙宽(Upper rostrum width,URW)、上侧壁长(UpperLateral wall length,ULWL)、下喙长(Lower Rostrum length,LRL)存在显著差异(P<0.01),同一性别不同群体角质颚的下头盖长(Lower Hood length,LHL)、下脊突长(Lower crest length,LCL)、LRL、下喙宽(Lower Rostrum width,LRW)、下侧壁长(LowerLateral wall length,LLWL)、下翼长(Lower Wing length,LWL)存在显著差异(P<0.01)。耳石形态参数经主成分分析,认为BNS群体雌雄的主成分因子主要集中在TSL/ML、DLL/ML、RW/ML和MW/ML,SPS群体主要集中于TSL/ML、RW/ML、WW/ML和DDL/ML;角质颚形态参数经主成分分析,认为BNS群体主成分因子主要集中在UHL/ML、UCL/ML、ULWL/ML和LRW/ML,SPS群体主要集中在UHL/ML、UCL/ML、ULWL/ML、URL/ML、LWL/ML和LRL/ML。利用角质颚和耳石对两群体样本分性别建立了判别函数,判别正确率均高于60%,所划分群体在部分形态指标上差异明显,具有一定的可信度。今后应加强样本采集个体大小和时间跨度,以更好的分析其群体变化规律。     

Root system architecture and gravity perception of a mangrove plant,Sonneratia alba J. Smith

We describe the features of the root system and the gravitropism of roots produced bySonneratia alba. The root system consists of four root types with different growth directions: (a) Pneumatophores, which are negatively orthogravitropic and their statocytes are very large (922 μm²) and the statolith is located near the proximal wall, (b) Cable roots and (c) Feeding roots which are both diagravitropic and their statoliths are settled along the longitudinal wall, and (d) Anchor roots which are positively orthogravitropic. The statocyte cells are the smallest (420 μm²) and statoliths settled at the distal wall. We found that all roots with marked gravitropism have statoliths that settle along different walls of the statocyte. This implies that the statoliths sensing of gravity is done by gravity on mass, and that they are denser than surrounding cytoplasm and this position is related to root growth direction. This finding matches the statoliths sediment under the effect of gravity. Irrespective of statolith, position and direction of growth may be stable.     

Growth and mating of southern African Lycoteuthis lorigera (Steenstrup, 1875) (Cephalopoda; Lycoteuthidae)

Lycoteuthis lorigera is an oceanic squid that is abundant in the Benguela system. Little is known about the biology of this squid except that it is eaten in large numbers by numerous oceanic predators and that males grow to larger size than females, which is unique for oegopsid squid. The aim of this study was to better understand the biology of this species by investigating its age and growth, as well as its mating system. Toward this end, the age of 110 individuals, ranging from 35 to 110 mm, was estimated by counting statolith growth increments. Estimates of age ranged from 131 to 315 days and varied with mantle length. No significant differences were found in the size of males and females of equivalent ages. The relationship between ML and age for both sexes was best described by an exponential growth curve, probably because no early life stages were aged in this study. Only one mature male (ML 160 mm) was aged, and preliminary estimates suggest it was 386 days old. Instantaneous growth rates were low (0.54% ML/day and 1.4% BM/day) but consistent with enoploteuthid growth rates. When the growth rate of L. lorigera was corrected for temperature encountered during the animal’s life, the growth rate was fast (0.47% BM/degree-days) and consistent with the hypothesis that small cephalopods grow fast and that large cephalopods grow older, rather than fast. Mature females were often mated and had spermatangia in a seminal receptacle on the dorsal pouch behind the nuchal cartilage. Males probably transfer spermatangia to the females using their long second and/or third arm pair since the paired terminal organs open far from the mantle opening. M. A. C. Roeleveld deceased.     

秘鲁外海茎柔鱼摄食洄游的稳定同位素研究

茎柔鱼是具有高度洄游性的中上层头足类,是生态系统中的关键物种,也是我国远洋渔业重要捕捞对象之一.了解掌握茎柔鱼摄食及洄游信息有助于对其资源进行更合理利用.头足类内壳是一良好的信息载体,其生长发育具有不可逆性且生长贯穿整个生活史过程,包含了头足类生活史过程中的全部信息.本研究通过对秘鲁外海茎柔鱼耳石日龄进行鉴定,建立其内壳叶轴生长方程,根据该生长方程对内壳叶轴进行连续切割,测定其碳、氮稳定同位素比值,并初步分析了其在生长发育过程中的食性转换和洄游习性.结果表明: 茎柔鱼个体在出生180 d后开始洄游活动,且营养层次存在下降趋势.证明了内壳连续取样分析茎柔鱼个体摄食习性和栖息地变化的可行性.     

西北印度洋海域鸢乌贼耳石微结构及生长特性

耳石是研究头足类年龄与生长、种群结构和生活史的重要硬组织之一。为研究西北印度洋鸢乌贼耳石微结构及生长特性,根据2019年2—5月我国灯光罩网渔船在西北印度洋调查生产期间采集的1009尾鸢乌贼样本,以耳石总长(TSL)、侧区长(LDL)、翼区长(WL)和最大宽度(MW)作为耳石的外形生长指标,结合日龄数据,对西北印度洋鸢乌贼耳石微结构及其生长特性进行了研究。结果表明: 根据生长纹间宽度,可将耳石微结构分为后核心区、暗区和外围区。协方差分析表明,TSL、LDL、WL和MW与日龄的关系均不存在显著的性别间差异,TSL、WL和MW与日龄的关系最适合用幂函数表示,而LDL与日龄的关系最适合用线性函数表示。总体而言,随着鸢乌贼日龄的增加,TSL、WL、LDL与MW的瞬时相对生长率与绝对生长率基本呈现下降的趋势, 140~180 d可能是西北印度洋鸢乌贼性成熟的日龄段。     

AGE AND GROWTH OF THE SQUID LOLIGO VULGARIS OFF THE SOUTH COAST OF PORTUGAL, USING STATOLITH ANALYSIS

Age composition and growth rates of the squid Loligo vulgaris(Lamark, 1797) were studied by examination of growth incrementswithin statohths of 419 specimens (mantle length, ML, rangingfrom 32 to 400 mm). The squid were obtained by monthly samplingfrom the catches of commercial trawls off southern Portugalbetween March and September, 1993 The total number of growthincrements in the mounted and ground statohths was counted usinga semi-automatic image analysis system. ML was significantlycorrelated with both the statolith length, TSL and the numberof increments, NI. The female statolith was slightly largerthan the male statolith for the same mantle size. Growth ratesof individuals showed high variability with an average estimatedat 34.6 mm month^–1 for males and 33.5 mm month^–1for females. Growth in length between 70 and 280 days was bestdescribed by a power function for both sexes. The growth indexof the statolith (TSL/NI) decreased with individual growth.This result may be related with the onset of sexual maturation.L. vulgaris hatched throughout the year with two distinct peaks,in spring which is the mam breeding period, and in autumn. Thelife cycle of the L. vulgaris population on the south Portugueseshelf was completed in one year ^*Present address for correspondence: Instituto de InvestigacionMariflas. Eduardo Cabello. 6 - 36208 VIGO. Spain (Received 28 November 1995; accepted 7 February 1996)     

Effects of pH on asexual reproduction and statolith formation of the scyphozoan, <Emphasis Type="Italic">Aurelia labiata</Emphasis>

Although anthropogenic influences such as global warming, overfishing, and eutrophication may contribute to jellyfish blooms, little is known about the effects of ocean acidification on jellyfish. Most medusae form statoliths of calcium sulfate hemihydrate that are components of their balance organs (statocysts). This study was designed to test the effects of pH (7.9, within the average current range, 7.5, expected by 2100, and 7.2, expected by 2300) combined with two temperatures (9 and 15°C) on asexual reproduction and statolith formation of the moon jellyfish, Aurelia labiata. Polyp survival was 100% after 122 d in seawater in all six temperature and pH combinations. Because few polyps at 9°C strobilated, and temperature effects on budding were consistent with published results, we did not analyze data from those three treatments further. At 15°C, there were no significant effects of pH on the numbers of ephyrae or buds produced per polyp or on the numbers of statoliths per statocyst; however, statolith size was significantly smaller in ephyrae released from polyps reared at low pH. Our results indicate that A. labiata polyps are quite tolerant of low pH, surviving and reproducing asexually even at the lowest tested pH; however, the effects of small statoliths on ephyra fitness are unknown. Future research on the behavior of ephyrae with small statoliths would further our understanding of how ocean acidification may affect jellyfish survival in nature.     

Abiotic influences on embryo growth: statoliths as experimental tools in the squid early life history

Statolith size and growth was used to determine the influence of abiotic factors on the growth of Loligo vulgaris and Sepioteuthis australis embryos. Recently spawned egg masses collected from the field were incubated in the laboratory under different levels of light intensity, photoperiod, or short periods of low salinity (30‰). Double tetracycline staining was used to follow statolith growth. In L. vulgaris constant light conditions produced significantly slower growth in the embryonic statoliths and embryos held at summer photoperiod had slower statolith growth than those held at winter photoperiods. However once they hatched out there was no evidence that photoperiod affected statolith growth. After hatching, in all photoperiods statolith growth rates decreased in comparison with late embryonic rates. In S. australis embryos, differences between the high and medium light intensities for summer and intermediate photoperiods were found, suggesting that under summer incubation temperature, longer daylengths at medium light intensity favoured higher statolith growth for this species. In comparison to controls, slower statolith growth in S. australis embryos due to low salinity only occurred when exposed for 72 h. Comparison with previous studies indicates that temperature seems to be the main abiotic factor influencing statolith growth during early stages, however, interactions among all abiotic factors needs to be determined as well as the unknown influence of other isolated factors, e.g., oxygen concentration within the egg mass.     

The structure of some cephalopod statoliths

Summary The statoliths of Sepia officinalis, Octopus vulgaris, Alloteuthis subulata and Taonius megalops have a smooth outline, but an irregular shape. They have projections and indentations. The statoliths from a pair of statocysts are usually quite similar in size and shape, and the general pattern is probably maintained throughout the size range of the species. Statoliths from large animals are marginally larger than those from smaller ones. The statolith usually occupies only a small part of the cavity of the statocyst, and it is situated in the anterior part of the statocyst. They are joined to the macula by hairs extending from it. These hairs are very delicate and easily broken during preparation of the specimens. The hairs are much longer and narrower than the receptor cilia of the macula. The receptor cilia are enclosed within holes in the tangled hairlike anchoring fibrils.The statolith is made up of crystalline subunits, the statoconia. The crystals vary in size, they are usually elongated, hexagonal with pointed ends. The statolith consists of a closely packed mass of these crystals, sometimes they are irregularly arranged, where in others they are stacked with their long axes parallel. In Sepia officinalis and Taonius megalops, the crystals are arranged in regular shaped packets and these packets of crystals are stacked together. These larger subunits are not always arranged in a regular way, and their major axes can be organised in several different ways. The size and outline of these large subunits do vary in different parts of the statolith.The external surface of the statolith is macroscopically smooth. Over some parts there is a surface layer covering the rod-like crystals that make up the major bulk of the stone. In other regions, the surface is rough at a microscopic level, the roughness is produced by the exposed ends of the filamentous crystals. The crystals are composed of calcium carbonate in the form of aragonite.I wish to thank Professor J.Z. Young, FRS, for considerable help, advice and encouragement throughout this study. Dr. A. Boyde generously allowed me to use his scanning electron microscope and gave freely of his expertise and time. Dr. J. Fitch kindly gave me some fossil statoliths and Dr. J. Elliott examined them with his x-ray diffraction apparatus. Dr. Marion Nixon helped me to collect and prepare the specimens. Mrs. E. Bailey, Miss P. Stephens and Mr. R. Moss provided the expert technical assistance