Geology topics

Nicholas S. Johnson

Publications and source records attributed to Nicholas S. Johnson.

At least 19 recordsLinked to original sources

USGS2026

DNA retention in sea lamprey digestive tracts: Insights from controlled feeding experiments

The sea lamprey ( Petromyzon marinus ), a non-native species in the Laurentian Great Lakes, has significantly impacted native fish communities and commercial fisheries, requiring population suppression efforts. While traditional control methods such as lampricides and barriers have reduced sea lamprey population abundance, questions remain regarding sea lamprey dietary composition given the focus of current damage assessments on economically and ecologically important host species. Recent advances in molecular technology offer promising methods of sea lamprey dietary assessment. Specifically, DNA metabarcoding enables species-specific identification of taxonomically diverse prey items from gut and fecal samples, and has proven effective in many taxa, including hematophagous species such as Arctic lamprey ( Lethenteron camtschaticum ) and sea lamprey. However, studies on DNA retention within digestive tracts are limited, particularly given the potential effects of environmental and dietary factors among hematophagous species. We used controlled feeding experiments to understand the effects these factors may have on DNA retention and host detectability within sea lamprey digestive tracts. Additionally, we evaluated the utility of metabarcoding for identifying multiple host species from consecutive feedings. Results indicate that host DNA can be detected up to 30 days post-feeding, with detection probability decreasing with increasing time following feeding. Temperature effects were dependent upon fasting periods, and host-switching trials indicated multiple previous host species could be detected from a single lamprey. Findings provide valuable insights for refining dietary analysis protocols for wild-caught sea lamprey within native and introduced ranges.

Environmental DNA↗

USGS2025

Ten lessons for controlling invasive species: Wisdom from the long-standing sea lamprey control program on the Laurentian Great Lakes

Sea lamprey ( Petromyzon marinus ) control in the Laurentian Great Lakes of North America is among the largest and most successful control programs of an invasive species anywhere on the planet. The effort began more than 75 years ago; it unites multiple nations, states, and provinces with the common goal of controlling this invasive species and protecting a valuable fishery. The science-based control program is administered by the Great Lakes Fishery Commission (GLFC), a body arising from a treaty signed by the United States and Canada. In the present article, we share 10 lessons learned from decades of successful sea lamprey control with the hopes of informing ongoing and future control programs targeting biological invasions. The 10 lessons we identified are to act boldly in times of crisis, to maintain the social license, to invest in capacity building, to break down the silos, to support fundamental science, to diversify your portfolio of control measures, to strive for continuous improvement, to confront the trade-off between information and action, to keep your foot on the gas, and to keep your eyes on the prize. The GLFC has long fostered a framework that uses some military strategy and verbiage that extends across the lessons (e.g., know your enemy). Other lessons are more nascent as the GLFC reenvisions its relationship with Indigenous peoples and governments in a path to reconciliation where two-eyed seeing is being embraced. Through adaptive management, horizon scanning methods, and embracing implementation science, the lessons learned about sea lamprey control will continue to evolve, which is itself a lesson. We submit that the lessons shared in the present article will help guide invasive species control programs spanning taxa, ecosystems, and regions.

Laurentian Great Lakes↗

USGS2025

Development of PCR blocking primers enabling DNA metabarcoding analysis of dietary composition in hematophagous sea lamprey

Conventional dietary assessments are challenging in hematophagous species, particularly in sea lamprey ( Petromyzon marinus ). However, recent technological developments and molecular approaches have provided an attractive alternative through the use of DNA metabarcoding. While DNA metabarcoding has been used for dietary analyses in numerous species, including lampreys, applications of universal primers that detect a diverse set of prey items can be limited by the amplification of predator DNA. In this study, we designed and tested eight blocking primers designed to suppress the amplification of sea lamprey DNA with vertebrate-universal primers targeting the mitochondrial 12S rRNA gene. This approach allowed for the use of a single marker to amplify a taxonomically diverse suite of host species, in contrast to previous studies that used multiple taxon-specific primer pairs (e.g., Salmonidae, Cyprinidae, and Catostomidae). Candidate blocking primers evaluated in this study differed in base pair length, end sequence modification, and purification method. Samples with different sea lamprey-to-host DNA ratios were subjected to multiple detection methods including gel electrophoresis, quantitative PCR, and DNA metabarcoding to assess the ability of each blocking primer to selectively suppress amplification of the sea lamprey 12S gene region. All blocking primers tested performed well and demonstrated high effectiveness, suppressing sea lamprey reads by > 99.9% in mock communities and improving host DNA sequence recovery across various sample types, including wild-caught lamprey. Results show that the blocking primers evaluated can facilitate molecular diet analysis in sea lamprey, allowing the amplification of a taxonomically diverse range of host fish species with universal primers.

Great Lakes↗

USGS2025

Field and laboratory evaluations of visible light as a cue for guiding downstream-migrating juvenile Sea Lamprey

Objective We evaluated white light as a potential guidance cue for juvenile Sea Lamprey Petromyzon marinus in a natural setting as well as the effect of water velocity (0.25-, 0.50-, 0.75-, and 1.0-m/s test velocities) on light guidance behavior in a controlled laboratory flume, and characterized emigration timing and movement rates in a small stream (∼10 m wide and 0.7 m deep). Methods Behaviors and rates of downstream movement were monitored using PIT telemetry in both studies. Results In the field study, downstream movement by juveniles released during October 30–November 27 appeared to be cued by precipitation-induced flow events when water temperatures ranged between 4°C and 8°C. Juveniles expressed lateral attraction to a short, bank-mounted linear light array, but the guidance effect was not strong or consistent between bank light locations. Downstream movement rates decreased slightly when juveniles were exposed to the light cue. In the laboratory flume experiment, at water velocities of 0.25 and 0.75 m/s, lamprey were 2.8 and 3.3 times more likely to be detected at antennas along a wall with a linear light array compared with other antennas across the width of the flume. Significant changes in distribution were detected farther upstream in the flume during 0.25- and 0.50-m/s water velocity trials compared with 0.75-m/s trials. Further, the rate of downstream movement through the length of the flume decreased under artificial lighting compared with dark controls under the 0.25- and 0.75-m/s velocity conditions. Conclusions The results suggest lamprey exhibit a behavioral response to the light cue in both lab and field, but water velocity influences how effectively juveniles can respond to light cues.

Massachusetts↗

USGS2025

Sea Lamprey control reduction during the COVID-19 pandemic corresponds to rapid increase in Sea Lamprey abundance

The Sea Lamprey Petromyzon marinus control program in the Laurentian Great Lakes is one of the longest-running and most successful invasive species suppression programs in the world. Although several techniques are used to suppress Sea Lamprey, the control program relies heavily on regular application of lampricide to kill stream-dwelling larvae. The COVID-19 pandemic disrupted lampricide application from 2020 to 2022, which provided a unique opportunity to test how Sea Lamprey populations and wound counts on fishes would respond. We evaluated the consequences of decreased control effort at a basin-wide level using standard control program metrics and through a focused analysis of multispecies wounding in Lake Ontario, which experienced the greatest disruption in Sea Lamprey control. The reduction in control effort corresponded to increased basin-wide adult Sea Lamprey abundance but was only weakly associated with Lake Trout Salvelinus namaycush wounding. However, the novel multispecies wounding rate metric calculated for Lake Ontario increased and was consistent with observations by anglers and managers regarding a sharp increase in Sea Lamprey abundance following reduced effort. Ultimately, our research highlights that consistent treatment is critically important for preventing damage stemming from Sea Lamprey and that reductions in suppression could quickly lead to a resurgence in abundance.

Great Lakes↗

USGS2025

Decision analysis of Integrated Pest Management: A case study on invasive sea lamprey in the Great Lakes Basin

Integrated Pest Management (IPM) provides a powerful framework for addressing threats to human well-being caused by nuisance species including invasives. We examined the hypothesis that adaptive management could erode barriers to IPM implementation by developing a decision-analytic adaptive management framework for invasive sea lamprey ( Petromyzon marinus ) IPM in the Laurentian Great Lakes of North America. The framework addressed objectives associated with coordinating multiple sea lamprey control actions at the regional scale and objectives associated with internal validity of control actions. We reduced the scope of possible management actions by orders of magnitude to the set of 6432 alternatives expected to be both socially acceptable and technically feasible. Using utility theory, we identified the management actions that optimized expected utility for all possible objective weighting schemes that considered tradeoffs between maximizing learning about control tactic efficacy and minimizing cost to the IPM program. Sensitivity analyses revealed that assumptions about the social acceptability of deploying electric weirs to control invasive sea lamprey influenced selection of the optimal control action, suggesting that resolving this source of uncertainty through iterative application of the framework may lead to improved sea lamprey control outcomes. Overall, we found that adaptive management enabled learning processes useful for overcoming barriers to IPM of invasive sea lamprey. It formalized learning about sea lamprey control tactic efficacy as an objective of the IPM institution, questioned previously held assumptions about what constitutes a viable control strategy, and enabled a management experiment with temporal and spatial replication.

Michigan, Ontario, Wisconsin↗

USGS2025

Larger larval sea lamprey (Petromyzon marinus) have longer survival times when exposed to the lampricide 3-trifluoromethyl-4-nitrophenol

Invasive sea lamprey ( Petromyzon marinus ) in the Laurentian Great Lakes have negatively impacted ecologically and economically important fishes for nearly a century. To mitigate these effects, the lampricide 3-trifluoromethyl-4-nitrophenol (TFM) is applied annually on a rotating basis to selected Great Lakes tributaries to kill larval lamprey before they become juveniles, out-migrate to the lakes, and parasitize other fishes. It has been hypothesized that larval size (e.g., mass, length) may affect survival time in response to TFM. To test this hypothesis, we conducted an experiment with 8611 larvae across four temporal replicates, in which TFM concentrations equivalent to those used in present-day stream treatments were applied for up to 18 h. When examining the survival times of larval lamprey exposed to TFM, we found a significant, positive relationship between length, mass, toxicity, and their interactions. For every 1 mm increase in total length, a corresponding increase by 1 g of mass reduced survival time by 0.4315 min [95 % CI: 0.5283–0.2992] and vice versa (i.e., the significant interaction between length and mass revealed that as larvae increase in mass, the survival benefit to being longer decreases, and vice versa). The changes in total length and mass of larval sea lamprey stored in ethanol for 4 months was also quantified. The observation that five larvae survived well past the 12-hour time window of a typical TFM field treatment highlights the need for continuous monitoring and the development of new control strategies to ensure the continued effective management of this invasive species.

Michigan↗

USGS2024

A decade-long study demonstrates that a population of invasive sea lamprey (Petromyzon marinus) can be controlled by introducing sterilized males

The release of sterilized insects to control pest populations has been used successfully during the past 6 decades, but application of the method in vertebrates has largely been overlooked or met with failure. Here, we demonstrate for the first time in fish, that a small population of sea lamprey ( Petromyzon marinus ; Class Agnatha), arguably one of the most impactful invasive fish in the world, can be controlled by the release of sterilized males. Specifically, the release of high numbers of sterile males (~ 1000's) into a geographically isolated population of adult sea lamprey resulted in the first multiyear delay in pesticide treatment since treatments began during 1966. Estimates of percent reduction in recruitment of age-1 sea lamprey due to sterile male release ranged from 7 to 99.9% with the precision of the estimate being low because of substantial year-to-year variability in larval density and distribution. Additional monitoring that accounts for recruitment variability in time and space would reduce uncertainty in the degree to which sterile male release reduces recruitment rates. The results are relevant to vertebrate pest control programs worldwide, especially as technical opportunities to sterilize vertebrates and manipulate sex ratios expand.

Michigan↗

USGS2024

Evaluation of larval sea lamprey Petromyzon marinus growth in the laboratory: Influence of temperature and diet

Conservation aquaculture provides a means for promoting environmental stewardship, useful both in the context of restoring native species and limiting the production of invasive species. Aquaculture of lampreys is a relatively recent endeavor aimed primarily at producing animals to support the restoration of declining native populations. However, in the Laurentian Great Lakes, where sea lamprey Petromyzon marinus are invasive, the ability to acquire a reliable source of certain life stages would be a significant benefit to those controlling their populations and studying the species. Here, we apply methodologies developed for Pacific lamprey Entosphenus tridentatus restoration to investigate the feasibility of rearing larval sea lamprey under laboratory conditions. In two experiments lasting 3 and 9 months, we tested the effects of different dietary sources and water temperature (ambient and controlled) on the survival and growth of wild-caught larvae. Rearing conditions had no effect on mortality, as larval survival was 100% in both experiments. Growth was significantly affected by water temperature, with the highest average daily growth rates observed at 22 and 15°C (0.14 mm day −1 ) and lowest at 8°C (0.06 mm day −1 ). Diets of yeast alone (0.19 and 0.21 g L −1 ) performed better than those comprising a mixture of yeast and other material when fed 3 times weekly (rice flour, wheat flour, fish meal; 0.19 and 0.32 g L −1 ). Averaged across the three constant temperatures (8, 15, and 22°C), larvae fed on yeast grew 0.13 mm day −1 and 0.01 g day −1 , whereas on yeast + fish meal, they grew 0.09 mm day −1 and 0.01 g day −1 . At ambient temperature (4–20°C), larvae fed on yeast grew 0.15 mm day −1 and 0.01 g day −1 , whereas those fed on yeast + wheat flour grew 0.13 mm day −1 and 0.008 g day −1 and those fed on yeast + rice flour grew 0.12 mm day −1 and 0.009 g day −1 . An experimental duration of 90 days was sufficient to detect significant changes to larval sea lamprey growth stemming from temperature variation. Overall, rearing of sea lamprey in captivity appears feasible at low density (31–32 g m −2 and 17–25 larvae m −2 ), but uncertainties remain regarding the most appropriate means of providing adequate feed for these fish in high-density conditions.

Aquaculture Research↗

USGS2024

Male lake char release taurocholic acid as part of a mating pheromone

The evolutionary origins of sexual preferences for chemical signals remain poorly understood, due, in part, to scant information on the molecules involved. In the current study, we identified a male pheromone in lake char ( Salvelinus namaycush ) to evaluate the hypothesis that it exploits a non-sexual preference for juvenile odour. In anadromous char species, the odour of stream-resident juveniles guides migratory adults into spawning streams. Lake char are also attracted to juvenile odour but have lost the anadromous phenotype and spawn on nearshore reefs, where juvenile odour does not persist long enough to act as a cue for spawning site selection by adults. Previous behavioural data raised the possibility that males release a pheromone that includes components of juvenile odour. Using metabolomics, we found that the most abundant molecule released by males was also released by juveniles but not females. Tandem mass spectrometry and nuclear magnetic resonance were used to identify the molecule as taurocholic acid (TCA), which was previously implicated as a component of juvenile odour. Additional chemical analyses revealed that males release TCA at high rates via their urine during the spawning season. Finally, picomolar concentrations of TCA attracted pre-spawning and spawning females but not males. Taken together, our results indicate that male lake char release TCA as a mating pheromone and support the hypothesis that the pheromone is a partial match of juvenile odour.

Journal of Experimental Biology↗

USGS2024

Protocol for monitoring and analyzing pheromone-mediated behavioral response of sea lamprey in a natural system

Olfactory-mediated behaviors in fish are often examined in artificial microcosms that enable well-controlled treatments but fail to replicate environmental and social contexts. However, observing these behaviors in nature poses challenges. Here, we describe a protocol for recording sea lamprey (Petromyzon marinus) behaviors in a natural system. We describe steps for administering and verifying accurate odorant concentrations, surveying sea lamprey abundance, and tracking sea lamprey movements. We also detail procedures to analyze treatment effects on pheromone-mediated spawning in a high-density population.

STAR Protocols↗

USGS2024

Evaluating behavioral responses of spawning-phase Walleyes to odors of rivers and other Walleyes

Many fish species show a remarkable ability to home to streams for spawning and fishery managers have applied that knowledge to restore or rehabilitate populations of some species. Walleye Sander vitreus show strong homing tendencies and the ability to select their natal river from other streams. The role of olfactory cues has been hypothesized but never tested in Walleye and many aspects of Walleye migratory behavior associated with spawning are poorly understood. The objectives of our study were: 1) to determine if spawning-phase Walleye from the Ford River in Michigan favored the odor of water from their home stream over that of a nearby river, and 2) to examine effects of male and female Walleye (conspecific) odors on water choice preferences of spawning-phase male Walleyes to learn whether such odors may attract adult Walleyes into rivers. Flow-through flume assays with two side-by-side channels were used to evaluate behavioral preferences of spawning-phase Walleye to river and conspecific odors. Odors of Ford River in 2021 trials and conspecific males in 2019 trials (when analyzed by individuals but not groups) influenced the behavior of male Walleye. However, follow up work during 2022 did not detect any responses to Ford River water or male-conditioned water. Additional work is needed to determine the possible role of olfactory cues in the homing behavior of spawning-phase Walleye.

Michigan↗

USGS2023

Evaluating the utility of effective breeding size estimates for monitoring sea lamprey spawning abundance

Sea lamprey ( Petromyzon marinus ) is an invasive species that is a significant source of mortality for populations of valued fish species across the North American Great Lakes. Large annual control programs are needed to reduce the species' impacts; however, the number of successfully spawning adults cannot currently be accurately assessed. In this study, effective breeding size ( N b ) and the minimum number of spawning adults ( N s ) were estimated for larval cohorts from 17 tributaries across all five Great Lakes using single nucleotide polymorphisms (SNP) genotyped via RAD-capture sequencing. Reconstructed larval pedigrees showed substantial variability in the size and number of full- and half-sibling groups, N b (<1–367), and N s (5–545) among streams. Generalized linear models examining the effects of stream environmental characteristics and aspects of sampling regimes on N b and N s estimates identified sample size, the number of sampling sites, and drainage area as important factors predicting N b and N s . Correlations between N b , N s , and capture–mark–recapture estimates of adult census size ( N c ) increased when streams with small sample sizes ( n < 50) were removed. Results collectively indicate that parameters estimated from genetic data can provide valuable information on spawning adults in a river system, especially if sampling regimes are standardized and physical stream covariates are included.

Ecology and Evolution↗

USGS2023

Synergistic behavioral antagonists of a sex pheromone reduce reproduction of invasive sea lamprey

Sex pheromones impart maximal attraction when their components are present at optimal ratios that confer balanced olfactory inputs in potential mates. Altering ratios or adding pheromone analogs to optimal mixtures may disrupt balanced olfactory antagonism and result in reduced attraction, however, tests in natural populations are lacking. We tested this hypothesis in sea lamprey ( Petromyzon marinus ), a fish whose male sex pheromone attracts females when two critical components, 3-keto petromyzonol sulfate (3kPZS) and petromyzonol sulfate (PZS), are present at certain ratios. Here, we report a pheromone analog, petromyzonol tetrasulfate (3sPZS), reduced female attraction to 3kPZS but not to PZS. 3sPZS mixed with additional PZS synergistically disrupted female attraction to the male pheromone and reduced spawning by 97% in a high-density population. Our results provide evidence of balanced olfactory antagonism in a vertebrate and establish a tactic to disrupt spawning of sea lamprey, a destructive invader of the Laurentian Great Lakes.

iScience↗

USGS2022

Pedigree analysis and estimates of effective breeding size characterize sea lamprey reproductive biology

The sea lamprey ( Petromyzon marinus ) is an invasive species in the Great Lakes and the focus of a large control and assessment program. Current assessment methods provide information on the census size of spawning adult sea lamprey in a small number of streams, but information characterizing reproductive success of spawning adults is rarely available. We used RAD-capture sequencing to genotype single nucleotide polymorphism (SNP) loci for ~1600 sea lamprey larvae collected from three streams in northern Michigan (Black Mallard, Pigeon, and Ocqueoc Rivers). Larval genotypes were used to reconstruct family pedigrees, which were combined with Gaussian mixture analyses to identify larval age classes for estimation of spawning population size. Two complementary estimates of effective breeding size ( N b ), as well as the extrapolated minimum number of spawners ( N s ), were also generated for each cohort. Reconstructed pedigrees highlighted inaccuracies of cohort assignments from traditionally used mixture analyses. However, combining genotype-based pedigree information with length-at-age assignment of cohort membership greatly improved cohort identification accuracy. Population estimates across all three streams sampled in this study indicate a small number of successfully spawning adults when barriers were in operation, implying that barriers limited adult spawning numbers but were not completely effective at blocking access to spawning habitats. Thus, the large numbers of larvae present in sampled systems were a poor indicator of spawning adult abundance. Overall, pedigree-based N b and N s estimates provide a promising and rapid assessment tool for sea lamprey and other species.

Michigan↗

USGS2022

Behavior of female adult Pacific lamprey (Entosphenus tridentatus) exposed to natural and synthesized odors

Conservation and management of Pacific Lamprey Entosphenus tridentatus and other imperiled lamprey species could include the use of chemosensory cues to attract or repel migrating adults. For restoration programs, passage of adult lamprey at dams might be improved by using cues to help guide lamprey through fishway entrances. In contrast, odors might repel unwanted invasive Sea Lamprey Petromyzon marinus in the Laurentian Great Lakes from spawning habitats or improve trapping efficiency. We conducted bioassays with Pacific Lamprey in a two-choice maze to evaluate the behavioral response of pre-ovulatory adult females to introduced chemical cues and changes in flow. During overnight tests, for each female we measured the number of entries into each arm of the maze and the amount of time spent in each arm after application of natural odors from pre-spawning conspecifics (males and females) in one of the arms. Using the same methods, we also tested whether adult females were attracted to natural odor from spermiating males, to a synthesized (artificially produced) component of a Sea Lamprey sex pheromone (3-keto petromyzonol sulfate, 3kPZS), or to an attraction flow (12 L/min as reference). In all tests, the lamprey showed consistent nocturnal activity, typically moving from sunset until sunrise and remaining inactive during daylight hours. For natural odors, the number of entries and the amount of time females spent in the treatment arm were not significantly different between control and treatment periods. However, females spent significantly less time in the treatment arm with the synthesized 3kPZS than when no odor was delivered. Females showed strong, positive responses to the attraction flow and with our assay, we could identify significant behavioral responses when the differences between the control and experimental means were greater than 15-20%. The response of lampreys to sex pheromones may be species-specific, with Pacific Lamprey less likely to respond to conspecific odors than Sea Lamprey.

Journal of Fish and Wildlife Management↗

USGS2022

Invasive species control and management: The sea lamprey story

Control of invasive species is a critical component of conservation biology given the catastrophic damage that they can cause to the ecosystems they invade. This is particularly evident with sea lamprey ( Petromyzon marinus ) in the Laurentian Great Lakes. Native to the Atlantic Ocean, the sea lamprey's ability to osmoregulate in fresh water, its wide thermal tolerance, generalist diet, and high fecundity allowed it to rapidly reach pest proportions in the prey-rich Great Lakes once it gained access through shipping canals. The invasion exacerbated declines in Great Lakes fisheries caused by overharvest, culminating in the crash of lake trout ( Salvelinus namaycush ) and other fish populations. In the last 60 years, however, a highly successful sea lamprey control program has reduced sea lamprey to ∼10% of their peak abundance and has been instrumental in enabling the rehabilitation of the Great Lakes ecosystem. In this chapter, we: (1) discuss the likely vectors of the invasion and the physiological attributes of sea lamprey that enabled them to become established in the Great Lakes; (2) review the two cornerstones of the sea lamprey control program—which relies on a combination of pesticides to eradicate multiple generations of larval sea lamprey in their nursey streams, and in-stream barriers to restrict the upstream migration of spawning lamprey—both of which exploit unique physiological vulnerabilities of sea lamprey; (3) describe how sea lamprey control can adversely affect non-target species and how these can be mitigated; (4) show how physiology-based approaches are improving our understanding of the lethal and sublethal effects of sea lamprey on host fishes; and (5) discuss the future of conservation physiology in sea lamprey control. The prime challenge in the next several decades of the Anthropocene will be to further refine the specificity of control tools while maintaining their efficacy, and to adapt to a warming climate and other anthropogenic activities affecting the Great Lakes and their tributaries.

Lake Erie, Lake Huron, Lake Michigan, Lake Ontario↗

USGS2021

Diel patterns of pheromone release by male sea lamprey

Costs to producing sexual signals can create selective pressures on males to invest signaling effort in particular contexts. When the benefits of signaling vary consistently across time, males can optimize signal investment to specific temporal contexts using biological rhythms. Sea lamprey, Petromyzon marinus , have a semelparous life history, are primarily nocturnal, and rely on pheromone communication for reproduction; however, whether male investment in pheromone transport and release matches increases in spawning activity remains unknown. By measuring (1) 3keto-petromyzonol sulfate (3kPZS, a main pheromone component) and its biosynthetic precursor PZS in holding water and tissue samples at six points over the course of 24 hours and (2) 3kPZS release over the course of several days, we demonstrate that 3kPZS release exhibits a consistent diel pattern across several days with elevated pheromone release just prior to sunset and at night. Trends in hepatic concentrations and circulatory transport of PZS and 3kPZS were relatively consistent with patterns of 3kPZS release and suggest the possibility of direct upregulation in pheromone transport and release rather than observed release patterns being solely a byproduct of increased behavioral activity. Our results suggest males evolved a signaling strategy that synchronizes elevated pheromone release with nocturnal increases in sea lamprey behavior. This may be imperative to ensure that male signaling effort is not wasted in a species having a single, reproductive event.

Integrative and Comparative Biology↗