Gill and intestinal Na+-K+ ATPase activity, and estimated maximal osmoregulatory costs, in three high-energy-demand teleosts: yellowfin tuna (Thunnus albacares), skipjack tuna (Katsuwonus pelamis), and dolphin fish (Coryphaena hippurus)

We hypothesize that the morpho-physiological adaptations that permit tunas to achieve maximum metabolic rates (MMR) that are more than double those of other active fishes should result in high water and ion flux rates across the gills and concomitant high osmoregulatory costs. The high standard metabolic rates (SMR) of tunas and dolphin fish may, therefore, be due to the elevated rates of energy expenditure for osmoregulation (i.e. teleosts capable of achieving exceptionally high MMR necessarily have SMR). Previous investigators have suggested a link between activity patterns and osmoregulatory costs based on Na⁺-K⁺ ATPase activity in the gills of active epipelagic and sluggish deep-sea fishes. Based on these observations, we conclude that high-energy-demand fishes (i.e. tunas and dolphin fish) should have exceptionally elevated gill and intestinal Na⁺-K⁺ ATPase activity reflecting their elevated rates of salt and water transfer. To test this idea and estimate osmoregulatory costs, we measured Na⁺-K⁺ ATPase activity (V _max) in homogenates of frozen samples taken from the gills and intestines of skipjack and yellowfin tunas, and the gills of dolphin fish. As a check of our procedures, we made similar measurements using tissues from hybrid red tilapia (Oreochromis mossambicus ×O. niloticus). Contrary to our supposition, we found no difference in Na⁺-K⁺ ATPase activity per unit mass of gill or intestine in these four species. We estimate the cost of osmoregulation to be at most 9% and 13% of the SMR in skipjack tuna and yellowfin tuna, respectively. Our results, therefore, do not support either of our original suppositions, and the cause(s) underlying the high SMR of tunas and dolphin fish remain unexplained. Received: 7 September 2000 / Accepted: 4 December 2000     

Behavior of bigeye (<Emphasis Type="Italic">Thunnus obesus</Emphasis>) and skipjack (<Emphasis Type="Italic">Katsuwonus pelamis</Emphasis>) tunas within aggregations associated with floating objects in the equatorial eastern Pacific

The horizontal and vertical movements of bigeye (Thunnus obesus Lowe, 1839) and skipjack (Katsuwonus pelamis Linnaeus, 1758) tunas within large multi-species aggregations associated with moored buoys or a drifting vessel were investigated, using ultrasonic telemetry and archival tags, along with sonar imaging, in the equatorial eastern Pacific Ocean (at 2°S–95°W and 2°N–95°W). Four sets of observations, each consisting of the concurrent monitoring of pairs of skipjack and/or bigeye with implanted acoustic or archival tags, were conducted in May 2002 and 2003. Ultrasonic telemetry data were not collected until 24 h or more after the fish were tagged and released, to avoid any abnormal behavior as a consequence of tagging. The pairs of acoustically tagged bigeye and skipjack, and also the entire aggregations, were primarily upcurrent of the moored buoy and downcurrent of the drifting vessel during the day. At night the aggregations were observed to be more diffuse, and the fish were feeding on organisms of the deep scattering layer. The aggregations returned to positions upcurrent of the buoy or downcurrent of the drifting vessel at dawn, commonly breezing at the surface within cohesive monospecific schools. The bigeye and skipjack had concurrent changes in depth records, occupying significantly greater mean depths at night than during the day, in most cases. When associated with a moored buoy, bigeye depth distributions were deeper during the day and night than those of skipjack, but bigeye depth distributions were shallower during the day and night than those of skipjack when associated with the drifting vessel. Simultaneous depth records of a large and a small bigeye with archival tags associated with a moored buoy also indicated diel changes in depth. The mean depth at night was significantly less than during the day for the larger bigeye, but the mean depth during the day was significantly less than during the night for the smaller bigeye. The mean depths during the day and night were significantly greater for the larger bigeye than the smaller.Electronic Supplementary Material  Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-004-1480-xCommunicated by J.P. Grassle, New Brunswick     

Allozyme and mitochondrial DNA variation in yellowfin tuna (Thunnus albacares) from the Pacific Ocean

Samples of yellowfin tuna (Thunnus albacares) collected in 1991 and 1992 from the western, central and eastern regions of the Pacific Ocean were examined for genetic variability. Four polymorphic allozyme loci (ADA ^*, FH ^*, GPI-S ^* and GPI-F ^*) were examined in all samples and a fifth polymorphism (GDA ^*) was examined in western and central samples only. Samples were also screened for mitochondrial DNA variation following restriction analysis by two enzymes (BcII and EcoRI) detecting polymorphic cut sites. Eighteen mtDNA haplotypes were revealed, with an overall nucleon diversity of 0.678. A subset of individuals screened for eight restriction enzymes had an overall nucleon diversity of 0.724 and a mean nucleotide diversity per sample of 0.359%. No significant spatial heterogeneity was detected for alleles at the ADA ^*, FH ^*, GPI-S ^* and GDA ^* loci nor for the mtDNA haplotypes. Significant heterogeneity was detected for GPI-F ^*. At this locus, the two eastern samples (southern California and northern Mexico) were not significantly different from each other but were significantly different (P<0.001) from the five western/central samples (Philippines, Coral Sea, Kiribati, Hawaii-91 and Hawaii-92). GPI-F ^* 100 was the most common allele in western and central regions, GPI-F ^* 75 the most common in eastern samples.     

Population structure of yellowfin tuna (Thunnus albacares) in the western Pacific Ocean, inferred from microsatellite loci

Five polymorphic microsatellite loci were examined in 1391 yellowfin tuna (Thunnus albacares) from eight regions of the western (Coral Sea, eastern Australia, Fiji, Indonesia, Philippines and Solomon Islands) and eastern (California and Mexico ) Pacific Ocean. Across all samples, numbers of alleles per locus ranged from 7 to 30 (mean: 17.0), and observed heterozygosities per locus ranged from 0.223 to 0.955 (mean: 0.593). Temporal collections were available for three areas: no significant temporal heterogeneity was observed for the Coral Sea (1991/1992 and 1995/1996 collections) or eastern Australia (1994/1995, 1995/1996, 1996/1997 and 1997/1998), but there was slight but significant heterogeneity at one locus (cmrTa-161) between the two Philippines collections (1994/1995 and 1996/1997). Genotypes generally showed a good fit to Hardy-Weinberg expectations within populations; only cmrTa-208 in the pooled Coral Sea population gave a significant deviation after Bonferroni correction for 40 tests, with a small but significant excess of homozygotes. Four loci showed no evidence of population differentiation following contingency Chi-squared and F_ST analyses. The fifth locus, cmrTa-161, showed small but significant differentiation (F_ST=0.002, P<0.001). This heterogeneity was largely a result of the Philippines 1994/1995 and Fiji collections; there was no correlation with geographic distance. The average F_ST across all five loci was very low (F_ST=0.002), but it was significant (P<0.001). It is unclear whether this low but significant differentiation reflects noise in the dataset, perhaps arising from experimental error, or real population differentiation. The finding of very limited population heterogeneity accords with most of the earlier allozyme and mitochondrial DNA studies of yellowfin tuna in the Pacific Ocean.     

Vertical movements, behavior, and habitat of bigeye tuna (Thunnus obesus) in the equatorial eastern Pacific Ocean, ascertained from archival tag data

The results presented in this report are based on analyses of 16,721 days of data downloaded from 96 archival tags recovered from bigeye tuna (Thunnus obesus; 54–159 cm in length, 0.97–5.44 years of age) at liberty from 31 to 1,508 days in the equatorial eastern Pacific Ocean. Analyses of daily timed depth and temperature records resulted in the classification of the data into three daily behavior types: characteristic, associative (associated with floating objects), and other. There is a significant positive correlation between the proportion of time fish exhibit characteristic behavior and increasing length, and significant negative correlations between the proportion of time bigeye exhibit associative and other behavior with increasing length. For the smallest (54–80 cm) to largest (100–159 cm) length classes, the vertical habitats utilized when exhibiting non-associative behaviors were 99 and 98% of the time above the thermocline depth (60 m) during the night, at the same average depth of 34 m, and 60 and 72% of the time below the thermocline during the day at average depths of 163 and 183 m, respectively. For the same smallest to largest length classes, when exhibiting associative behavior, the average nighttime and daytime depths were 25 and 21, and 33 and 37 m, respectively. The apparent effects of the environment on the behavior of the fish are discussed.     

Behavior of yellowfin (<Emphasis Type="Italic">Thunnus albacares</Emphasis>) and bigeye (<Emphasis Type="Italic">T. obesus</Emphasis>) tuna in a network of fish aggregating devices (FADs)

The influence of multiple anchored fish aggregating devices (FADs) on the spatial behavior of yellowfin (Thunnus albacares) and bigeye tuna (T. obesus) was investigated by equipping all thirteen FADs surrounding the island of Oahu (HI, USA) with automated sonic receivers (“listening stations”) and intra-peritoneally implanting individually coded acoustic transmitters in 45 yellowfin and 12 bigeye tuna. Thus, the FAD network became a multi-element passive observatory of the residence and movement characteristics of tuna within the array. Yellowfin tuna were detected within the FAD array for up to 150 days, while bigeye tuna were only observed up to a maximum of 10 days after tagging. Only eight yellowfin tuna (out of 45) and one bigeye tuna (out of 12) visited FADs other than their FAD of release. Those nine fish tended to visit nearest neighboring FADs and, in general, spent more time at their FAD of release than at the others. Fish visiting the same FAD several times or visiting other FADs tended to stay longer in the FAD network. A majority of tagged fish exhibited some synchronicity when departing the FADs but not all tagged fish departed a FAD at the same time: small groups of tagged fish left together while others remained. We hypothesize that tuna (at an individual or collective level) consider local conditions around any given FAD to be representative of the environment on a larger scale (e.g., the entire island) and when those conditions become unfavorable the tuna move to a completely different area. Thus, while the anchored FADs surrounding the island of Oahu might concentrate fish and make them more vulnerable to fishing, at a meso-scale they might not entrain fish longer than if there were no (or very few) FADs in the area. At the existing FAD density, the ‘island effect’ is more likely to be responsible for the general presence of fish around the island than the FADs. We recommend further investigation of this hypothesis.

---