The DOI matches its Crossref record; 32 of the 33 references it cites were found in a registry or an index; none of the 29 sources its claims cite is checked for support yet.
The energy costs of wading in water
Open accessAbstract
Studies measuring the energy costs of wading in water have been limited to higher walking speeds in straight lines, in deep water. However, much foraging in water, by both humans and other primates, is conducted in the shallows and at low speeds of locomotion that include elements of turning, as befits searching for cryptic or hidden foods within a patch. The present study brings together data on the rate of oxygen consumption during wading by humans from previous studies, and augments these with new data for wading in shallower depths, with slower and more tortuous walking, to obtain a better understanding both of the absolute costs of wading in typical scenarios of aquatic foraging and of how the cost of wading varies as a function of water depth and speed of locomotion. Previous and present data indicate that, at low speeds, wading has a similar energetic cost to walking on land, particularly at lower water depths, and only at higher speeds is the cost of wading noticeably more expensive than when water is absent. This is probably explained by the relatively small volume of water that must be displaced during locomotion in shallow waters coupled with the compensating support to the limbs that the water affords. The support to the limbs/body provided by water is discussed further, in the context of bipedal locomotion by non-human primates during wading.
Overview
AI snapshot
Written from the paper’s full text.
TL;DR
In shallow water at slow, tortuous walking speeds, human wading costs about the same energy as walking on land; only at higher speeds does wading become noticeably more expensive. The study combines new Douglas-bag measurements in knee- and thigh-high water with reanalyzed published data to argue that typical aquatic foraging up to waist-deep carries little extra energetic cost.
Key findings
- New data: mass-specific rate of oxygen consumption during wading in knee-high (N=10) and thigh-high (N=10) water was similar to walking on land across the tested slow speeds; one-way ANOVAs at four speeds all returned p values above 0.15.
- Effect sizes were typically small, particularly between land walking and thigh-high wading; knee-high wading was always slightly lower than the other conditions except at the highest speed.
- Combined with previous data, wading in waist- or chest-high water also converges with land walking below around 0.3 m s−1; only above approximately 0.35 m s−1 is energy expenditure noticeably higher than on land.
- The authors suggest water’s support to the limbs/body may offset the cost of displacing water in shallows, with implications for bipedal wading in non-human primates.
Methods
- Participants walked on land and in water around a circular path (2.4 m diameter pool, 0.8 m high, effective capacity about 2700 L) at cadences of 33, 45, 60 and 90 steps min−1, with an additional 75 steps min−1 for wading; they took small steps and performed 180° turns. Wading conditions were knee height (N=10) or thigh height (N=10), with several participants undertaking both; land control data are shown for N=16.
- Oxygen consumption was measured using the Douglas bag method; participants walked at each cadence for at least two minutes before measurement, then expired air was collected for a further minute or more. Surface water temperature varied between 15.4°C and 17.5°C.
- Analysis used one-way ANOVAs comparing the three conditions at each of four speeds after checking normality and homogeneity of variance. Published data were compiled by digitizing figures from Kuliukas et al. and Ghesquiere and Bunkens, benchmarking relative values against land-walking estimates, and estimating mean body mass of 65 kg for Ghesquiere and Bunkens.
Limitations
- The paper does not state limitations explicitly.
- Evident from the text: small samples (N=10 per wading depth; land control N=16), several participants undertook both experimental conditions, control data at 0.41 m s−1 were unavailable, and water temperature was limited to 15.4–17.5°C.
- Evident from the text: previous data were extracted via Plot Digitizer and Ghesquiere and Bunkens’ mean body mass was estimated at 65 kg; the protocol used short steps and turns, so it may not generalize to all wading, and no non-human primate metabolic data were collected.
Model-written summary. Check it against the paper before relying on it.
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Claims the paper makes
0 of 29 cited sources checked · 0 supported · 0 flagged
- Findingown data
In the present study, the effect sizes (differences in mean mass-specific rate of oxygen consumption between conditions at each speed) are typically quite small, particularly between the control condition and the experimental condition where water was at thigh height.
“It can be seen that the effect sizes (the differences in mean mass-specific between the conditions at each speed) are typically quite small, particularly between the control condition and the experimental condition where water was at thigh height”
Quote in the paperSubstantial claim - Findingown data
Typical wading behaviour, i.e. in shallow water (knee- or thigh-high water) and at slow walking speeds, incurs a similar cost of transport to walking on land at the same speeds.
“The effect sizes in the present study show that typical wading behaviour, i.e. in shallow water (knee- or thigh-high water) and at slow walking speeds, incurs a similar cost of transport to walking on land at the same speeds (Fig. 1).”
- Also stated in: § Abstract · hedged
“Previous and present data indicate that, at low speeds, wading has a similar energetic cost to walking on land, particularly at lower water depths”
Quote in the paperSubstantial claim - Also stated in: § Abstract · hedged
- Premisecitation
Low speeds and frequent changes in direction associated with foraging will likely serve to increase transport costs.
“Low speeds and also frequent changes in direction associated with foraging will likely serve to increase transport costs (Halsey and White, 2012; Wilson et al., 2013).”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Wading at shallower depths attenuates the cooling effects of water.
“Furthermore, wading at shallower depths attenuates the cooling effects of water (M. Lieger and C. Niemtiz, unpublished data)”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites M. Lieger and C. Niemtiz, unpublished data
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Premisecitation
Kuliukas et al. provide evidence that the energy cost of bipedal walking is reduced in water, albeit in humans undertaking bent-knee bent-hip walking.
“Indeed, Kuliukas et al. provide evidence of such, albeit in humans undertaking bent-knee bent-hip walking (Kuliukas et al., 2009).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Kuliukas et al., 2009 source
- Premisenone given
Only two papers have been published on the energy costs of wading through water.
“However, only two papers have been published on the energy costs of wading through water.”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Ghesquiere and Bunkens, 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Kuliukas et al., 2009 source
- Premisecitation
Previous studies measuring the energy costs of wading in water have been limited to higher walking speeds in straight lines, in deep water.
“Studies measuring the energy costs of wading in water have been limited to higher walking speeds in straight lines, in deep water.”
- Also stated in: § INTRODUCTION · plain
“The two previous studies on the energy costs of wading have measured straight line walking”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Ghesquiere and Bunkens, 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- cites Kuliukas et al., 2009 source
- Also stated in: § INTRODUCTION · plain
- Premisecitation
The data of Kuliukas et al. are reported as factorial values relative to a single transport cost and thus cannot be easily compared to the values reported by Ghesquiere and Bunkens.
“These data are reported as factorial values relative to a single transport cost and thus cannot be easily compared to the values reported by Ghesquiere and Bunkens (Ghesquiere and Bunkens, 1991).”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Kuliukas et al., 2009 source
- cites Ghesquiere and Bunkens, 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
- Interpretationreasoning
Typical wading by humans during foraging in water up to waist-deep does not incur noteworthy energy costs over and above those incurred during terrestrial foraging.
“Therefore, it might be reasonable to conclude that typical wading by humans during foraging in water up to waist-deep does not incur noteworthy energy costs over and above those incurred during terrestrial foraging.”
Quote in the paperSubstantial claim - Interpretationcitation
The increased costs in the present study are certainly in part due to the inclusion of turns in the walking protocol, which incur a significant energetic cost.
“However, these increased costs will certainly be in part due to the inclusion of turns in the walking protocol, which incur a significant energetic cost (Wilson et al., 2013).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Wilson et al., 2013 source
- Premisecitation
All of the many other primate species observed wading typically exhibit an orthograde (hence bipedal) posture, rising up onto their hind limbs.
“Humans are not the only primate species known to wade and interestingly, all of the many other primate species observed wading typically exhibit an orthograde (hence bipedal) posture, rising up onto their hind limbs (western lowland gorillas (Doran and McNeilage, 1998), e.g. chimpanzees (Karlowski, 1996), bonobos (Myers Thompson, 2002), several species of monkey (Niemitz, 2010)).”
Quote in the paperNot a substantial claim0 of 3 sources checked - Premisecitation
Much foraging in water, by both humans and other primates, is conducted in the shallows and at low speeds of locomotion that include elements of turning, as befits searching for cryptic or hidden foods within a patch.
“However, much foraging in water, by both humans and other primates, is conducted in the shallows and at low speeds of locomotion that include elements of turning, as befits searching for cryptic or hidden foods within a patch.”
Quote in the paperSubstantial claim0 of 2 sources checked - Premisecitation
Wading in water is considered to provide significant support to a quadrupedal animal in an orthograde posture, even in the shallows.
“Wading in water is considered to provide significant support to a quadrupedal animal in an orthograde posture, even in the shallows (Niemitz, 2010).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Niemitz, 2010 source
- Premisecitation
There is little evidence from metabolic data that humans, or an early hominin Australopithecus afarensis, experience(d) low energy expenditure during pedestrian locomotion in comparison to other mammals in general.
“however, in fact, there is little evidence from metabolic data that humans, or an early hominin Australopithecus afarensis, experience(d) low energy expenditure during pedestrian locomotion in comparison to other mammals in general (Halsey and White, 2012; Rubenson et al., 2007).”
Quote in the paperSubstantial claim0 of 2 sources checked
- It may state two separate claims as one. 9
- Probably not a substantial claim. 8
It may state two separate claims as one.
FindingcitationPeople walking in water up to their waists and up to their chests also experience similarly low costs of transport at slower wading speeds; even with such water depths, only at walking speeds above approximately 0.35 m s−1 is rate of energy expenditure noticeably higher than that on land.
“Indeed, people walking in water up to their waists (Kuliukas et al., 2009) and up to their chests (Ghesquiere and Bunkens, 1991) also experience similarly low costs of transport at slower wading speeds; even with such water depths, only at walking speeds above ∼0.35 m”
Quote in the paperSubstantial claim0 of 2 sources checked- cites Kuliukas et al., 2009 source
- cites Ghesquiere and Bunkens, 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim. The paper’s conclusions may not depend on it. It may state two separate claims as one.
Findingown dataThe standard errors for each condition mean are small, although larger at the highest speed, suggesting that the sample values are typically fairly accurate estimates.
“The standard errors for each condition mean are small (Fig. 1), although larger at the highest speed, suggesting that the sample values are typically fairly accurate estimates.”
Quote in the paperNot a substantial claimIt may state two separate claims as one.
FindingreasoningFor a given locomotion speed, mass-specific rate of oxygen consumption is somewhat higher in the present study than in both of the previous studies, even though the wading conditions in the present study involved lower water depths.
“for a given locomotion speed is somewhat higher in the present study than in both of the previous studies (Fig. 2), even though the wading conditions in the former involved lower water depths.”
Quote in the paperNot a substantial claimIt may state two separate claims as one.
Findingown dataOne-way ANOVAs comparing the three conditions at each of four speeds yielded all p values above 0.15, providing little evidence for true differences in mass-specific rate of oxygen consumption between the conditions at each speed.
“All resultant p values were above 0.15 and thus provided little evidence for true differences in mass-specific between the conditions at each of the speeds.”
Quote in the paperSubstantial claimIt may state two separate claims as one.
InterpretationreasoningThe present data provide more realistic absolute values of immediate energy expenditure for people foraging for cryptic or hidden food, whether in water or on land, and confirm the interpretation that the added cost of walking in water is at most minimal at low speeds.
“Thus arguably the present data provide more realistic absolute values of immediate energy expenditure for people foraging for cryptic or hidden food, whether in water or on land, and certainly confirm our interpretation of the data from the previous studies that the added cost of walking in water is at most minimal at low speeds.”
Quote in the paperSubstantial claimProbably not a substantial claim. It may state two separate claims as one.
PremisecitationTerrestrial bipedal walking, although possible by non-human primates for short periods, may be energetically expensive for them.
“there have been a number of arguments put forward that as such, terrestrial bipedal walking, although possible by these species for short periods of time, may be energetically expensive (Carey and Crompton, 2005; Leonard and Robertson, 2001; Nakatsukasa et al., 2004; Pontzer et al., 2009; Rodman and McHenry, 1980).”
Quote in the paperSubstantial claim0 of 3 sources checked- cites Carey and Crompton, 2005 source
- cites Leonard and Robertson, 2001 source
- cites Nakatsukasa et al., 2004 source
- Rests on an assumption: All non-human primates are obligate quadrupeds.
It may state two separate claims as one.
InterpretationcitationThe higher mass-specific rate of oxygen consumption in the present study is perhaps mainly explained by the short step lengths exhibited by the participants to simulate foraging behaviour, which may result in the feet being in contact with the ground for a shorter duration per step for a given speed, and locomotion costs are thought to increase with the inverse of foot contact time.
“This is perhaps mainly explained by the short step lengths exhibited by the participants to simulate foraging behaviour. Shorter step lengths may result in the feet being in contact with the ground for a shorter duration per step for a given speed, and locomotion costs are thought to increase with the inverse of foot contact time (Kram and Taylor, 1990).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Kram and Taylor, 1990 source
Probably not a substantial claim.
Premisenone givenAs yet there is no information on the costs of wading in water at thigh or knee height.
“Furthermore, as yet there is no information on the costs of wading in water at thigh or knee height.”
Quote in the paperSubstantial claimProbably not a substantial claim. The paper’s conclusions may not depend on it.
InterpretationcitationAn alternative explanation for the higher mass-specific rate of oxygen consumption in the present study is that shorter steps result in an attenuation of the pendular exchange of kinetic and potential energy exhibited in the normal, striding human gait.
“An alternative explanation is that shorter steps result in an attenuation of the pendular exchange of kinetic and potential energy exhibited in the normal, striding human gait (Cavagna et al., 1977).”
Quote in the paperNot a substantial claim0 of 1 sources checked- cites Cavagna et al., 1977 source
Probably not a substantial claim.
PremisecitationKuliukas et al. reported data for wading in various depths of water, speeds and with varying knee flexion.
“Kuliukas et al. reported data for wading in various depths of water, speeds and with varying knee flexion (Kuliukas et al., 2009).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Kuliukas et al., 2009 source
Probably not a substantial claim. The paper’s conclusions may not depend on it. The paper may give no reason why it holds. It may state two separate claims as one.
InterpretationreasoningTrying to locomote through shoulder-deep water typically triggers swimming behaviour in humans, probably at least in part because of the effort needed to move through the water column due to the volume of water that must be displaced and also the loss of traction on the ground due to the positive buoyancy of the body.
“while for humans at least, trying to locomote through shoulder-deep water typically triggers swimming behaviour, probably at least in part because of the effort needed to move through the water column due to the volume of water that must be displaced and also the loss of traction on the ground due to the positive buoyancy of the body.”
Quote in the paperNot a substantial claimIt may state two separate claims as one.
PremisecitationIn reality wading probably tends to consist of small steps and some degree of turning to remain within a foraging patch and respond to cues about food location.
“while in reality wading probably tends to consist of (a) small steps as both a response to moving through a more viscous medium and the advantage that slower locomotion affords when searching within foraging patches (Viswanathan et al., 1999), (b) some degree of turning to remain within a foraging patch and respond to cues about food location.”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Viswanathan et al., 1999 source
Probably not a substantial claim.
PremisecitationGhesquiere and Bunkens reported a large increase in rate of energy expenditure in water compared to on land, particularly at higher walking speeds, where the water reached up to the chest (armpits) of the participants.
“Ghesquiere and Bunkens reported a large increase in rate of energy expenditure in water compared to on land, particularly at higher walking speeds, where the water reached up to the chest (armpits) of the participants (Ghesquiere and Bunkens, 1991).”
Quote in the paperSubstantial claim0 of 1 sources checked- cites Ghesquiere and Bunkens, 1991
Couldn’t be matched to a paper in Crossref, so it can’t be checked.
Probably not a substantial claim.
InterpretationreasoningIf at least some quadrupedal primates do indeed incur an energy cost during bipedal walking, then it is reasonable to suppose that this cost is attenuated while in water due to the support of that medium.
“if at least some quadrupedal primates do indeed incur an energy cost during bipedal walking then it is reasonable to suppose that this cost is attenuated while in water due to the support of that medium.”
Quote in the paperSubstantial claim- Rests on an assumption: All non-human primates are obligate quadrupeds.
The paper relies on this without support.
All non-human primates are obligate quadrupeds.
“All non-human primates are obligate quadrupeds”
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Self-citation
Flagged first, then by citations
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Metadata
- Authors
- Halsey, L. G., Tyler, C. J., & Kuliukas, A. V.
- Affiliations
- University of Roehampton, Holybourne Avenue, London SW15 4JD, UK
- University of Western Australia, 35 Stirling Highway, Crawley, WA 6009, Australia
- Venue
- Biology Open
- Publisher
- The Company of Biologists
- Published
- 2014-06-06
- Volume
- 3
- Issue
- 7
- Pages
- 571-574
- ISSN
- 2046-6390 (ISSN-L)
- PMC
- PMC4154292
- OpenAlex
- W2130452794
- Cited by
- 5
- References
- 33
- Citations by year
- 2025: 1 · 2023: 1 · 2021: 1 · 2019: 1 · 2016: 1
- FWCI
- 0.28
- Type
- journal article
- Language
- en
- Open access
- gold
- doi.org
- Locations
- Biology Open (publishedVersion, cc-by)
- Europe PMC (PubMed Central) (publishedVersion, cc-by)
- PubMed Central (publishedVersion, cc-by)
- UWA Profiles and Research Repository (UWA) (publishedVersion, other-oa)
- PubMed (publishedVersion)
- DOAJ (DOAJ: Directory of Open Access Journals) (submittedVersion)
- HighWire Press Open Archive (submittedVersion)
- Topics
- Life Sciences › Agricultural and Biological Sciences › Ecology, Evolution, Behavior and Systematics › Primate Behavior and Ecology
- Physical Sciences › Environmental Science › Ecology › Physiological and biochemical adaptations
- Health Sciences › Medicine › Physiology (medical) › Cardiovascular and exercise physiology
- Keywords
- wading, energy costs, oxygen consumption, water depth, locomotion speed, bipedalism, non-human primates, limb support, water displacement
About these checks
Registry, retraction, Cites and statistics results are computed, not judged, with one exception: a printed reference that no index lists is read once by a model before it counts as a citation error. Claims are extracted by a model and kept only when their quote appears verbatim in the text; each cited source is then checked, with a model’s help, for whether it says what the claim relies on. Those verdicts can be wrong — read the claim’s quote and the cited source before relying on one.
A language model drew the claims from the paper on 2026-11-10, under version 3 of the claim definition (model deepseek-v4.1-flash, prompt extract-v3).
A second model labelled each claim on 2026-11-10 (model jev-1.13-20260917, definition revision gold-guideline-v0.2).
Statistics and arithmetic are recomputed, not judged.
Confidence bands are shown only for fields calibrated on our annotated test set. None is calibrated yet, so no band is shown.
What counts as a claim ›How we measure confidence ›How we check this ›
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