How much energy does that mouthful cost?

By Lisa Hildebrand, PhD student, OSU Department of Fisheries, Wildlife, & Conservation Sciences, Geospatial Ecology of Marine Megafauna Lab

Tagging a whale is no easy feat, nor is it without some impact to the whale – no matter how minimized through the use of non-penetrating suction cup tags. Yet, in August 2021 the GEMM Lab initiated a new phase in our research on gray whales, aimed at obtaining a better understanding of the underwater lives and energetics of a gray whale (Figure 1, top image). We captured some amazing data through these specialized, non-invasive tags that provide a brief window into their world and physiology. The video recordings from the tags showed us whales digging their heads into the benthos generating billowing clouds of sediment, likely exploiting desirable prey patches (Figure 1, middle images). We also saw foraging whales undertake dizzying spins and headstands for hours, demonstrating the fascinating maneuverability and flexibility of gray whales (Figure 1, bottom image). But what is motivating us to capture this information?

The GEMM Lab has researched the ecology and physiology of Pacific Coast Feeding Group (PCFG) gray whales since 2015. Our efforts have filled crucial knowledge gaps to better understand this sub-group of the Eastern North Pacific (ENP) gray whale population. We now know that gray whale body condition increases throughout a foraging season and can fluctuate considerably between years (Soledade Lemos et al. 2020). Additionally, body condition varies significantly by reproductive state, with calves and pregnant females displaying higher body conditions (Soledade Lemos et al. 2020). We have also validated and quantified fecal steroid and thyroid hormone metabolite concentrations, providing us with thresholds to identify a stressed vs. a not stressed whale based on its hormone levels (Lemos et al. 2020). These validations have allowed us to make correlations between poor body condition and the steroid hormone cortisol which confirm that slim whales are stressed, while chubby whales are relaxed (Lemos et al. 2021). These physiological results are particularly salient in the light of our recent findings that PCFG gray whales select prey quality over prey quantity when foraging (Hildebrand et al. in review) and that the caloric content of available prey species in the PCFG range vary significantly (Hildebrand et al. 2021).

While we have addressed several fundamental questions about the PCFG in the last 7 years, answering one question has led to asking 10 more questions – a common pattern in science. Given that we know (1) PCFG whales improve their body condition over the course of the foraging season (Soledade Lemos et al. 2020), (2) PCFG females are able to successfully give birth to and wean calves (Calambokidis & Perez 2017), and (3) certain prey in the PCFG region are of higher caloric value than prey in the ENP Arctic foraging grounds (Hildebrand et al. 2021), a big question that we continue to scratch our heads about is why does the PCFG sub-group have such a small abundance (~250 individuals; Calambokidis et al. 2017) in comparison to the much larger ENP population (~21,000 individuals; Stewart & Weller 2021). Several hypotheses have been suggested including that the energetic costs of feeding may differ between ENP and PCFG whales, with the latter having to expend more energy to obtain prey due to the different foraging behaviors employed (Torres et al. 2018) to obtain diverse prey types, thus justifying the larger abundance of the ENP (Hildebrand et al. 2021). 

Quantifying the energetic cost of baleen whale behaviors is not simple. However, the development of animal-borne tags has allowed scientists to make big strides regarding behavioral cost quantification. The majority of this work has focused on rorqual whales (i.e., blue, humpback, fin whales; e.g., Goldbogen et al. 2013; Cade et al. 2016) as their characteristic lunge-feeding strategy produces a distinct signal in the accelerometer sensors integrated within the tags, making feeding events easier to identify. Gray whales, unlike rorquals, do not lunge-feed. ENP gray whales predominantly feed benthically; diving down to the benthos where they turn onto their side and suction mouthfuls of soft sediment (mud) that contains amphipods that they filter out of the mud (Nerini & Oliver 1983). PCFG whales feed benthically as well, but they also use a number of other feeding behaviors to obtain a variety of prey in a variety of benthic habitats, including headstands, bubble blasts, and sharking (Torres et al. 2018). The above-mentioned gray whale feeding behaviors involve much subtler movements than the powerful, distinctive lunges displayed by rorquals, yet they undoubtedly still incur some energetic cost to the whales. However, exactly how energetically costly the various gray whale feeding behaviors are remains unknown.

One of the three suction cup tags we deployed on gray whales. Dr. Cade printed special “kelp shields” (blue part of the tag) to prevent kelp from potentially getting caught underneath the tag since PCFG whales often forage on reefs with a lot of kelp. This tag includes a video camera (the lens can be seen in the center of the tag) to record video of the whale’s underwater behavior. Source: L. Torres.

This knowledge gap is one of the reasons why the GEMM Lab initiated a new project in close collaboration with Dr. Dave Cade from Stanford University and John Calambokidis from Cascadia Research Collective to quantify and understand the energetics and underwater behavior of gray whales using suction-cup tags. The project was kick-started with a very successful pilot effort the week of August 16th this year. Tags were placed on the backs of three different PCFG gray whales with a long carbon fiber pole and attached to the whales with four suction cups. The tags recorded video, position, accelerometry, and magnetometry data, which we will use to recreate the animal’s movements (pitch, roll), heading, trackline, and environment. Although the weather forecast did not look promising for most of the week, we lucked out with perfect conditions for one day during which we managed to deploy three tags on three different gray whales that are well-known, long-term study animals of the GEMM Lab. The tags stayed on the whales for 1-6 hours and were all recovered (including an adventurous trip up the Alsea River which involved a kayak deployment!). 

Dr. Cade spent the rest of the week teaching GEMM Lab PI Leigh Torres, University of British Columbia Master’s student Kate Colson (who is co-advised by Leigh and Dr. Andrew Trites), and myself the intricacies of data download, processing, and preliminary analysis of the tag data. For her Master’s research, Kate will develop a bioenergetics model for the PCFG sub-group that includes data on foraging energetics (estimated from the tag data) and prey availability in the PCFG foraging range. I plan on using the tag data to assess behavior patterns of PCFG whales relative to habitat as part of my PhD research. All together analysis of the data from these short-term tag deployments will help us get closer to understanding the behavioral choices, habitat needs, and energetic trade-offs of whales living in a rapidly changing ocean. With the success of this pilot effort, we plan to conduct another suction-cup tagging effort next summer to hopefully capture and explore more mysterious underwater behaviors of the PCFG.

An ecstatic team at the end of a very long yet successful day of suction cup tagging. Bottom (from left): Leigh Torres, Lisa Hildebrand, Clara Bird, Dave Cade, KC Bierlich. Top: John Calambokidis.

This project was funded by sales and renewals of the special Oregon whale license plate, which benefits MMI. We gratefully thank all the gray whale license plate holders, who made this research effort possible.

Literature cited

Cade, D. E., Friedlaender, A. S., Calambokidis, J., & Goldbogen, J. A. 2016. Kinematic diversity in rorqual whale feeding mechanisms. Current Biology 26(19):2617-2624. doi:10.1016/j.cub.2016.07.037.

Calambokidis, J., & Perez, A. 2017. Sightings and follow-up of mothers and calves in the PCFG and implications for internal recruitment. IWC Report SC/A17/GW/04 for the Workshop on the Status of North Pacific Gray Whales (La Jolla: IWC). 

Calambokidis, J., Laake, J., & Perez, A. 2017. Updated analysis of abundance and population structure of seasonal gray whales in the Pacific Northwest, 1996-2015. IWC Report SC/A17/GW/05 for the Workshop on the Status of North Pacific Gray Whales (La Jolla: IWC).

Goldbogen, J. A., Friedlaender, A. S., Calambokidis, J., McKenna, M. F., Simon, M., & Nowacek, D. P. 2013. Integrative approaches to the study of baleen whale diving behavior, feeding performance, and foraging ecology. BioScience 63(2):90-100. doi:10.1525/bio.2013.63.2.5.

Hildebrand, L., Bernard, K. S., & Torres, L. G. 2021. Do gray whales count calories? Comparing energetic values of gray whale prey across two different feeding grounds in the eastern North Pacific. Frontiers in Marine Science 1008. doi:10.3389/fmars.2021.683634.

Lemos, L. S., Olsen, A., Smith, A., Burnett, J. D., Chandler, T. E., Larson, S., Hunt, K. E., & Torres, L. G. 2021. Stressed and slim or relaxed and chubby? A simultaneous assessment of gray whale body condition and hormone variability. Marine Mammal Science. doi:10.111/mms.12877.

Lemos, L.S., Olsen, A., Smith, A., Chandler, T.E., Larson, S., Hunt, K., and L.G. Torres. 2020. Assessment of fecal steroid and thyroid hormone metabolites in eastern North Pacific gray whales. Conservation Physiology 8:coaa110.

Nerini, M. K., & Oliver, J. S. 1983. Gray whales and the structure of the Bering Sea benthos. Oecologia 59:224-225. doi:10.1007/bf00378840.

Soledade Lemos, L., Burnett, J. D., Chandler, T. E., Sumich, J. L., & Torres, L. G. 2020. Intra- and inter-annual variation in gray whale body condition on a foraging ground. Ecosphere 11(4):e03094.

Stewart, J. D., & Weller, D. W. 2021. Abundance of eastern North Pacific gray whales 2019/2020. Department of Commerce, NOAA Technical Memorandum NMFS-SWFSC-639. United States: NOAA. doi:10.25923/bmam-pe91.

Torres, L.G., Nieukirk, S.L., Lemos, L., and T.E. Chandler. 2018. Drone Up! Quantifying Whale Behavior From a New Perspective Improves Observational Capacity. Frontiers in Marine Science:


Whale blow: good for more than spotting whales

Clara Bird, Masters Student, OSU Department of Fisheries and Wildlife, Geospatial Ecology of Marine Megafauna Lab

Whale blow, the puff of air mixed with moisture that a whale releases when it comes to the surface, is a famously thrilling indicator of the presence of a whale. From shore, spotting whale blow brings the excitement of knowing that there are whales nearby. During boat-based field work, seeing or hearing blow brings the rush of adrenaline meaning that it’s game time. Whale blow can also be used to identify different species of whales, for example gray whale blow is heart shaped (Figure 1). However, whale blow can be used for more than just spotting and identifying whales. We can use the time between blows to study energetics.

Figure 1. Gray whale blow is often heart shaped (when there is very little wind). Source:

A blow interval is the time between consecutive blows when a whale is at the surface (Stelle, Megill, and Kinzel 2008). These are also known as short breath holds, whereas long breath holds are times between surfacings (Sumich 1983).  Sumich (1983) hypothesized that short breath holds lead to efficient rates of oxygen use. The body uses oxygen to create energy, so “efficient rate of oxygen use” means that longer breath holds do not use much more oxygen and subsequently do not produce more energy.  Surfacings, during which short blow intervals occur, are often thought of as recovery periods for whales. Think of it this way, when you sprint, immediately afterwards you typically need to take a break to just breathe and recover.

We hypothesize that we can use blow intervals as a measure of how strenuous an activity is; shorter blow intervals may indicate that an activity is more energetically demanding (Wursig, Wells, and Croll 1986). Let’s go back to the sprinting analogy and compare the energetic demands of walking and running. Imagine I asked you to walk for five minutes, stop and measure the time between each breath, and then run for five minutes and do the same; after running, you would likely breathe more heavily and take more breaths with less time between them. This result indicates that running is more demanding, which we already know because we can do other experiments with humans to study metabolic rate and related metrics. In the case of gray whales, we cannot do experiments in the same way, but we can use the same analogy. Several studies have examined how blow intervals differ between travelling and foraging.

Wursig, Wells, and Croll (1986) measured blow interval, surfacing time, and estimated dive depth and duration of gray whales in Alaska from a boat during the foraging season. They found that blow intervals were shorter during feeding. They also found that the number of blows per surfacing increased with increasing depth. Overall these findings suggest that during the foraging season, feeding is more strenuous than other behaviors and that deeper dives may be more physiologically stressful.

Stelle, Megill, and Kinzel (2008) studied gray whales foraging off of British Columbia, Canada. They found shorter blow intervals during foraging, intermediate blow intervals during searching, and longer blow intervals during travelling. Interestingly, within feeding behaviors, they found a difference between whales feeding on mysids (krill-like animals that swim in the water column) and whales feeding benthically on amphipods. They found that whales feeding on mysids made more frequent but shorter dives with short blow intervals at surface, while whales feeding benthically had longer dives with longer blow intervals. They hypothesized that this difference in surfacing pattern is because mysids might scatter when disturbed, so gray whales surface more often to allow the mysids swarm to reform. These studies inspired me to start investigating these same questions with my drone video data.

As I review the drone footage and code the behaviors I also mark the time of each blow. I’ve done some initial video coding and using this data I have started to look into differences in blow intervals. As it turns out, we see a similar difference in blow interval relative to behavior state in our data: whales that are foraging have shorter blow intervals than when traveling (Figure 2). It is encouraging to see that our data shows similar patterns.

Figure 2. Boxplot of mean blow interval per sighting of foraging whales and travelling whales.

Next, I would like to examine how blow intervals differ between foraging tactics. A significant part of my thesis is dedicated to studying specific foraging tactics. The perspective from the drone allows us to identify behaviors in greater detail than studies from shore or boat (Torres et al. 2018), allowing us to dig into the differences between the different foraging behaviors. The purpose of foraging is to gain energy. However, this gain is a net gain. To understand the different energetic “values” of each tactic we need to understand the cost of each behavior, i.e. how much energy is required to perform the behavior. Given previous studies, maybe blow intervals could help us measure this cost or at least compare the energetic demands of the behaviors relative to each other. Furthermore, because different behaviors are likely associated with different prey types (Dunham and Duffus 2001), we also need to understand the different energetic gains of each prey type (this is something that Lisa is studying right now, check out the COZI project to learn more). By understanding both of these components – the gains and costs – we can understand the energetic tradeoffs of the different foraging tactics.

Another interesting component to this energetic balance is a whale’s health and body condition. If a whale is in poor health, can it afford the energetic costs of certain behaviors? If whales in poor body condition engage in different behavior patterns than whales in good body condition, are these patterns explained by the energetic costs of the different foraging behaviors? All together this line of investigation is leading to an understanding of why a whale may choose to use different foraging behaviors in different situations. We may never get the full picture; however, I find it really exciting that something as simple and non-invasive as measuring the time between breaths can contribute such a valuable data stream to this project.


Dunham, Jason S., and David A. Duffus. 2001. “Foraging Patterns of Gray Whales in Central Clayoquot Sound, British Columbia, Canada.” Marine Ecology Progress Series 223 (November): 299–310.

Stelle, Lei Lani, William M. Megill, and Michelle R. Kinzel. 2008. “Activity Budget and Diving Behavior of Gray Whales (Eschrichtius Robustus) in Feeding Grounds off Coastal British Columbia.” Marine Mammal Science 24 (3): 462–78.

Sumich, James L. 1983. “Swimming Velocities, Breathing Patterns, and Estimated Costs of Locomotion in Migrating Gray Whales, Eschrichtius Robustus.” Canadian Journal of Zoology 61 (3): 647–52.

Torres, Leigh G., Sharon L. Nieukirk, Leila Lemos, and Todd E. Chandler. 2018. “Drone up! Quantifying Whale Behavior from a New Perspective Improves Observational Capacity.” Frontiers in Marine Science 5 (SEP).

Wursig, B., R. S. Wells, and D. A. Croll. 1986. “Behavior of Gray Whales Summering near St. Lawrence Island, Bering Sea.” Canadian Journal of Zoology 64 (3): 611–21.