The impact of overfishing and El Niño on the condition factor and reproductive success of Peruvian hake, Merluccius gayi peruanus

In this study we examine physiological responses of Peruvian hake (Merluccius gayi peruanus) to changes in their population structure and physical environment during the period 1971–2004. In particular, we assess the relative investment of energy in growth and/or reproduction of small (20–35 cm) and large (35–50 cm) hake. We calculated the (i) condition (Fulton’s K), (ii) gonad and (iii) gut fullness indices for 42,761 female hakes sampled from commercial landings; these indices indicate fish somatic, reproductive and feeding condition, respectively. Using Generalized Additive Models we then examined potential relationships between these indices and sea surface temperature anomalies and date. Drastic energy exhaustion and a decrease in female hake fecundity were observed during El Niño events. The long-term trend showed a general increase in condition factor and a decrease in gonad index for large hake between 1971 and 2004. Small hake exhibited a different trend with an increase in reproductive activity, which was accompanied by an earlier maturation. We hypothesise that the observed low investment of energy in reproduction by large female hake might be related to the lack of large males, due to a sex-selective fishery and the impact of El Niño. We suggest that fishing diminished hake reproductive capacity, modified the sex ratio in favour of females and increased population vulnerability to environmental stress, in particular to the El Niño. The impact of multidecadal variability and predators like the squid, Dosidicus gigas, remain unresolved until longer time series become available.     

驼背鲈引种驯养及人工诱导性腺发育和繁殖

为探讨通过人工诱导获得驼背鲈成熟亲鱼的途径,在室内水池进行驼背鲈驯养研究;用口服和注射性激素、控温、控光、营养强化、模拟生态条件等方法处理驼背鲈以诱导其性腺发育。结果表明,控光和模拟生态条件不能有效诱导性腺发育;激素、控温和营养强化相结合可诱导驼背鲈性腺发育并成熟。对其中１ 组诱导成熟的亲鱼（雌、雄各１ 尾）进行人工催产后,亲鱼自然产卵受精,孵出仔鱼２ 万尾。驼背鲈在驯养条件下生长较快;通过人工控制和诱导可达到性腺发育和成熟,性腺发育速度较快。     

海湾扇贝性腺发育的初步研究

１９９２年７月－１１月在青岛胶州五号码头养殖海区采集海湾扇贝，定期取样，测壳长、水温，并对性腺进行切片观察，以研究其性腺发育。观察表明，海湾扇贝性腺由３部分构成：滤泡、生殖管和生殖输送管。     

全人工配合饵料投喂越冬亲虾试验

通过两年度的小试和生产性试验,从越冬亲虾入池至产卵过程全部投喂人工配合饵料,其成活率和性腺发育要好于投沙蚕、四角蛤的鲜活饵料组(对照组),生产性试验的产卵量、孵化率和幼体变态率优于对照组。试验用饵的成本比对照组低45.3％。     

黑斑口虾蛄(Oratosquilla Kempi Schmitt)配子发生的显微观察

观察黑斑口虾蛄Oratosquillakempi的精巢和卵巢 ,并对精、卵巢进行组织切片显微观察。结果表明 :黑斑口虾蛄的精巢由 1对很细的管状结构组成 ,精子发育可分为精原细胞、初级精母细胞、次级精母细胞、精子等期 ,精子圆球形 ,无鞭毛。卵巢分左、右两叶 ,对称且相互粘连。卵细胞为多黄卵 ,卵母细胞发育分为未发育期、卵黄形成前期、卵黄形成期和成熟期 4期 ;在卵黄形成前期形成大的生发泡。卵巢发育分为未发育期、发育早期、发育期、成熟前期、成熟期和产卵后期 6期。     

环沟格特哈(Katelysia rimularis)的生殖周期

周年采集环沟格特蛤（Katelysiarimularis）分别测定生长指标（总重、软体部重、性腺重、肝重和软体部干重等）并用组织切片观察性腺发育的状况，以确定性腺发育的周期和时间。组织的光镜研究说明环沟格特蛤雌雄性腺发育均可分为增殖期、生长期、成熟期、排放期和休止期。在厦门吴冠镇环沟格特蛤1月开始其性腺发育,3月进入生长期，5～6月成熟，5～11月都有产卵，每年7月11月各有一次排卵高峰期。     

黄河三角洲河蚬(Corbicula fluminea)的繁殖生物学研究

自2009年2月—2010年1月,共采集黄河三角洲流域分布的河蚬498个,采用组织学方法分析其性腺的年周期变化,利用考马斯亮蓝方法进行了蛋白质含量的年周期测量。结果表明,黄河三角洲分布的河蚬雌雄异体,性比为1:1,性腺发育分为6个时期:增殖期、生长期、成熟期、排放期、恢复期和排尽休止期。配子发生期开始于2月份,一年内有两个繁殖高峰期,分别为每年的6月中旬和10月上旬。蛋白质含量无性别差异,闭壳肌和外套膜的蛋白质含量全年都处于较低水平,无明显波动,性腺-内脏团和足的蛋白质含量相对较高,并分别在5月份和5、8月份达到峰值,与河蚬的繁殖盛期显著相关。     

Environmental drivers of small scale spatial variation in the reproductive schedule of a commercially important bivalve mollusc

Understanding variability in reproductive schedules is essential to the management of recruitment limited fisheries such as that of Pecten maximus. Small scale (<5 km) variation in gonad condition and the onset of spawning of P. maximus were found among commercial scallop grounds in Isle of Man waters. Environmental and fishing drivers of these spatial patterns were investigated using a generalised additive model. Rate of change in temperature over the month prior to sampling was identified as the short term driver of gonad weight associated with the autumn spawning event. Long term drivers were average annual chlorophyll a concentration, scallop density, stratification index and shell size. The model explained 42.8% of deviance in gonad weight. Within site variation in gonad condition was high, indicating a “bet hedging” reproductive strategy which may decrease the chance of fertilisation especially at low densities. Therefore, areas protected from fishing, where scallop densities can increase may help buffer against reproductive failure. An increase in shell length from 100 mm to 110 mm equated to an increase of approximately 20% in gonad weight. Protecting scallops from fishing mortality until 110 mm (age four) compared to 100 mm (age three) may lead to an overall increase in lifetime reproductive output by a factor of 3.4.     

Zooplankton communities and oceanographic structures in a high-resolution grid in the south-eastern corner of the Bay of Biscay

A one year long monthly sampling series at a fixed station was combined with a high-spatial resolution survey (May 2003) to investigate factors controlling zooplankton distribution in the south-eastern corner of the Bay of Biscay. Species abundance and gonad maturity for Calanoides carinatus and Calanus helgolandicus were estimated to clarify whether the observed spatial patterns were attributable to increased population growth or to mechanical accumulation. The two studies included distinct oceanographic regimes. The eastern corner of the grid was characterized by Adour river plume waters and the Cap Breton canyon, and the rest of the area by an alternating pattern of physical convergences and divergences. In the river plume, the zooplankton community diversity and abundances were highest in response to the nutrient enriched river plume waters and the submarine canyon. In the rest of the area, distribution and abundance patterns were the result of a combination of behaviour and physical transport.     

Life history strategies in zooplankton communities: The significance of female gonad morphology and maturation types for the reproductive biology of marine calanoid copepods

The present review addresses the reproductive strategies of marine calanoid copepods, as affected by their physiological preconditioning, and aims to enhance understanding of their adaptations to specific environmental conditions.Knowledge about oocyte development and internal gonad structure, especially in relation to feeding conditions, is essential for a complete understanding of the reproductive strategies of the copepods. Therefore, the foci of the review are to identify general patterns in oocyte and gonad development in calanoid copepod species from marine ecosystems worldwide and to elucidate the significance of gonad structures for reproductive strategies.Oogenesis is similar in all copepod species. During maturation, the morphology of the oocytes changes distinctly and, according to oocyte size and appearance of ooplasm and nucleus, five oocyte developmental stages are distinguished. In contrast, the gonad structure and its changes during the spawning cycle differ considerably among species, and these differences are related to specific reproductive traits.Four gonad morphology types can be distinguished: the Calanus-type, found in species from all over the world with distinctly different life history traits, is apparently most common in calanoid copepods. In this gonad type, most oocyte developmental stages are present simultaneously, and usually many oocytes mature synchronously, all of which are released in one clutch. The gonad structure allows frequent spawning and large clutches, hence, high egg production rates. This may be a preconditioning for exploiting seasonally high food supply. However, the Calanus-type was also found in species producing eggs at lower rates. In the diverticula of Pseudocalanus-type gonads, only two oocyte developmental stages are present and usually fewer oocytes mature synchronously. Accordingly, the egg production rate is generally lower as compared to the Calanus-type, and apparently only this gonad-type is structurally suitable for ovigerity. Species with Pseudocalanus-type gonads are present from polar seas to the tropics, some of them being key species. The Acartia-type was scarce, found in only one species, Acartia clausi. Here all oocyte developmental stages are present, including intermediate stages, but only a few oocytes mature synchronously and are released together. High spawning frequency compensates for the small clutches, and hence egg production rate may be as high as in Calanus-type gonads. In the Aetidius-type gonad, the total number of oocytes in the diverticula is low as is the number of oocytes maturing synchronously. Less is known about the reproductive biology of species with Aetidius-type gonads; however, their distribution and feeding patterns suggest that this type is common in species inhabiting environments of low food availability.The differences in gonad structures also lead to differences in the egg size:female size ratio, as the space available for each mature oocyte depends on the total number of oocytes. Independent from gonad-type, the eggs are relatively large in species in which the gonads contain only few oocytes, whereas small eggs are produced by species with gonads filled with many oocytes. Since all species carrying their eggs in external sacs until hatching (ovigerous species) have Pseudocalanus-type gonads, the scatter in their egg size:female size ratio is low. The broadcast spawning species are of all gonad-types, and consequently the scatter among them is high.A major factor affecting the timing and magnitude of spawning of calanoid copepods is the energy supply for gonad development. Therefore, part of the review elucidates the role of internal and external resources in fuelling egg production. In many species, freshly assimilated food is transferred into egg material within a short period of time, and clutch size and spawning frequency are the two parameters that allow adjustment of egg production to food availability and temperature. However, internal body reserves may also fuel oocyte development. The extent to which oogenesis depends on ambient feeding conditions varies considerably among species and even within species, apparently reflecting their adaptation to specific environmental conditions. All copepod species continue to release eggs for a short period after the onset of starvation, indicating that a short-term nutrient pool fuels final oocyte maturation. Some species, especially from polar and subpolar regions, are capable of reproducing on internal reserves, either solely or in addition to freshly ingested food, hence, decoupling the reproduction to some extent from the spring bloom. The reproductive strategy of Calanus finmarchicus, which responds with extreme flexibility to variations in food supply, is presented here as an example for the complexity of the reproductive processes in a calanoid copepod.In accordance with their physiological pre-conditioning, the calanoid copepod species exhibit distinctly different reproductive patterns. Polar and sub-polar environments are characterized by strong seasonality in light regime, and hence, phytoplankton concentration. The reproductive activity of many dominant (herbivorous) species peaks during the spring bloom. However, species that utilize internal reserves or food resources other than phytoplankton, reproduce before and after the bloom, or even year-round. In the North Atlantic, presented here as an example for a boreal marine ecosystem, maximum reproduction of the dominant copepod species Calanus finmarchicus occurs during the spring bloom. However, the timing of peak spawning varies between areas, as the onset of the bloom varies with hydrography and climate. The subtropical Gulf of Aqaba, in contrast to other subtropical systems, is characterized by seasonal phytoplankton succession driven by thermal winter convection. These changing environmental conditions, however, did not affect the reproduction of two of the dominant species, Clausocalanus farrani and C. furcatus. In contrast, other calanoid species exhibit reproductive cycles, either related to temperature (Ctenocalanus vanus) or to ontogenetic migration (Rhincalanus nasutus).The information gained from studies of morphology and reproductive traits contributes to standardization of methods in reproductive studies. Based on detailed knowledge of gonad morphology and its changes during maturation and due to food supply, classification schemata have been developed allowing the identification of females ready to spawn. In a next step, egg production may be estimated from preserved females by assessing clutch size through the number of maturing oocytes in the gonads. This approach, however, is as yet applicable only to the species best studied, Calanus finmarchicus.The present review shows that morphological studies on gonad maturation processes may provide significant contributions to fundamental ecological questions. Thus, they extend our knowledge of reproductive ecology from simply relating reproductive traits to abiotic and biotic factors toward a mechanistic understanding of how reproduction is regulated in calanoid copepods.