Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design
Identifieur interne : 001776 ( Istex/Corpus ); précédent : 001775; suivant : 001777Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design
Auteurs : D V Lauritzen ; F S Hertel ; L K Jordan ; M S GordonSource :
- Bioinspiration & Biomimetics [ 1748-3182 ] ; 2010.
Abstract
Behavioral and kinematic properties and capacities of wild migratory salmonid fishes swimming upstream and jumping up waterfalls generally have played only minor roles in the design and construction of passageways intended to help these fishes get past dams and other human-made obstacles blocking their movements. This paper reports the results of an experimental study of relevant behavioral and kinematic properties of adult kokanee salmon (Oncorhynchus nerka) jumping up waterfalls as they migrate upstream. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. Kokanee salmon spontaneously jump up waterfalls within a relatively narrow range of conditions, including low flow speeds, near vertical angles and pool depth to fall height ratios near 1.0. Preferred values for each parameter are, to some extent, dependent on other parameters. In contrast to previous misconceptions, jumping behavior is initiated by running S-start accelerations from beneath the boils formed in the plunge pools below waterfalls, as opposed to C-start standing jumps from the surface. S-starts are immediately followed by burst swimming to the point of takeoff at the surface. These results can contribute to an improved basis for developing designs of fish passageways that may ultimately make them more effective and efficient.
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DOI: 10.1088/1748-3182/5/3/035006
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<record><TEI wicri:istexFullTextTei="biblStruct"><teiHeader><fileDesc><titleStmt><title xml:lang="en">Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design</title><author><name sortKey="Lauritzen, D V" sort="Lauritzen, D V" uniqKey="Lauritzen D" first="D V" last="Lauritzen">D V Lauritzen</name><affiliation><mods:affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: dlauritz@ccsf.edu</mods:affiliation></affiliation></author><author><name sortKey="Hertel, F S" sort="Hertel, F S" uniqKey="Hertel F" first="F S" last="Hertel">F S Hertel</name><affiliation><mods:affiliation>Department of Biology, California State UniversityNorthridge, Northridge, CA 91330, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: fritz.hertel@csun.edu</mods:affiliation></affiliation></author><author><name sortKey="Jordan, L K" sort="Jordan, L K" uniqKey="Jordan L" first="L K" last="Jordan">L K Jordan</name><affiliation><mods:affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: msgordon@ucla.edu</mods:affiliation></affiliation></author><author><name sortKey="Gordon, M S" sort="Gordon, M S" uniqKey="Gordon M" first="M S" last="Gordon">M S Gordon</name><affiliation><mods:affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: ljordan@ucla.edu</mods:affiliation></affiliation></author></titleStmt><publicationStmt><idno type="wicri:source">ISTEX</idno><idno type="RBID">ISTEX:9ACE70A18185FE1AA829B2846A52A448DEC8EDA0</idno><date when="2010" year="2010">2010</date><idno type="doi">10.1088/1748-3182/5/3/035006</idno><idno type="url">https://api.istex.fr/document/9ACE70A18185FE1AA829B2846A52A448DEC8EDA0/fulltext/pdf</idno><idno type="wicri:Area/Istex/Corpus">001776</idno><idno type="wicri:explorRef" wicri:stream="Istex" wicri:step="Corpus" wicri:corpus="ISTEX">001776</idno></publicationStmt><sourceDesc><biblStruct><analytic><title level="a" type="main" xml:lang="en">Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design</title><author><name sortKey="Lauritzen, D V" sort="Lauritzen, D V" uniqKey="Lauritzen D" first="D V" last="Lauritzen">D V Lauritzen</name><affiliation><mods:affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: dlauritz@ccsf.edu</mods:affiliation></affiliation></author><author><name sortKey="Hertel, F S" sort="Hertel, F S" uniqKey="Hertel F" first="F S" last="Hertel">F S Hertel</name><affiliation><mods:affiliation>Department of Biology, California State UniversityNorthridge, Northridge, CA 91330, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: fritz.hertel@csun.edu</mods:affiliation></affiliation></author><author><name sortKey="Jordan, L K" sort="Jordan, L K" uniqKey="Jordan L" first="L K" last="Jordan">L K Jordan</name><affiliation><mods:affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: msgordon@ucla.edu</mods:affiliation></affiliation></author><author><name sortKey="Gordon, M S" sort="Gordon, M S" uniqKey="Gordon M" first="M S" last="Gordon">M S Gordon</name><affiliation><mods:affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</mods:affiliation></affiliation><affiliation><mods:affiliation>E-mail: ljordan@ucla.edu</mods:affiliation></affiliation></author></analytic><monogr></monogr><series><title level="j">Bioinspiration & Biomimetics</title><title level="j" type="abbrev">Bioinsp. Biomim.</title><idno type="ISSN">1748-3182</idno><imprint><publisher>IOP Publishing</publisher><date type="published" when="2010">2010</date><biblScope unit="volume">5</biblScope><biblScope unit="issue">3</biblScope><biblScope unit="page" from="1">1</biblScope><biblScope unit="page" to="11">11</biblScope><biblScope unit="production">Printed in the UK</biblScope></imprint><idno type="ISSN">1748-3182</idno></series><idno type="istex">9ACE70A18185FE1AA829B2846A52A448DEC8EDA0</idno><idno type="DOI">10.1088/1748-3182/5/3/035006</idno><idno type="PII">S1748-3182(10)52493-1</idno><idno type="articleID">352493</idno><idno type="articleNumber">035006</idno></biblStruct></sourceDesc><seriesStmt><idno type="ISSN">1748-3182</idno></seriesStmt></fileDesc><profileDesc><textClass></textClass><langUsage><language ident="en">en</language></langUsage></profileDesc></teiHeader><front><div type="abstract">Behavioral and kinematic properties and capacities of wild migratory salmonid fishes swimming upstream and jumping up waterfalls generally have played only minor roles in the design and construction of passageways intended to help these fishes get past dams and other human-made obstacles blocking their movements. This paper reports the results of an experimental study of relevant behavioral and kinematic properties of adult kokanee salmon (Oncorhynchus nerka) jumping up waterfalls as they migrate upstream. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. Kokanee salmon spontaneously jump up waterfalls within a relatively narrow range of conditions, including low flow speeds, near vertical angles and pool depth to fall height ratios near 1.0. Preferred values for each parameter are, to some extent, dependent on other parameters. In contrast to previous misconceptions, jumping behavior is initiated by running S-start accelerations from beneath the boils formed in the plunge pools below waterfalls, as opposed to C-start standing jumps from the surface. S-starts are immediately followed by burst swimming to the point of takeoff at the surface. These results can contribute to an improved basis for developing designs of fish passageways that may ultimately make them more effective and efficient.</div></front></TEI><istex><corpusName>iop</corpusName><author><json:item><name>D V Lauritzen</name><affiliations><json:string>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</json:string><json:string>E-mail: dlauritz@ccsf.edu</json:string></affiliations></json:item><json:item><name>F S Hertel</name><affiliations><json:string>Department of Biology, California State UniversityNorthridge, Northridge, CA 91330, USA</json:string><json:string>E-mail: fritz.hertel@csun.edu</json:string></affiliations></json:item><json:item><name>L K Jordan</name><affiliations><json:string>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</json:string><json:string>E-mail: msgordon@ucla.edu</json:string></affiliations></json:item><json:item><name>M S Gordon</name><affiliations><json:string>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</json:string><json:string>E-mail: ljordan@ucla.edu</json:string></affiliations></json:item></author><subject><json:item><lang><json:string>eng</json:string></lang><value>salmon</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>kokanee</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>jumping</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>waterfalls</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>behavior</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>kinematics</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>fish passageways</value></json:item><json:item><lang><json:string>eng</json:string></lang><value>restoration ecology</value></json:item></subject><language><json:string>eng</json:string></language><originalGenre><json:string>paper</json:string></originalGenre><abstract>Behavioral and kinematic properties and capacities of wild migratory salmonid fishes swimming upstream and jumping up waterfalls generally have played only minor roles in the design and construction of passageways intended to help these fishes get past dams and other human-made obstacles blocking their movements. This paper reports the results of an experimental study of relevant behavioral and kinematic properties of adult kokanee salmon (Oncorhynchus nerka) jumping up waterfalls as they migrate upstream. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. Kokanee salmon spontaneously jump up waterfalls within a relatively narrow range of conditions, including low flow speeds, near vertical angles and pool depth to fall height ratios near 1.0. Preferred values for each parameter are, to some extent, dependent on other parameters. In contrast to previous misconceptions, jumping behavior is initiated by running S-start accelerations from beneath the boils formed in the plunge pools below waterfalls, as opposed to C-start standing jumps from the surface. S-starts are immediately followed by burst swimming to the point of takeoff at the surface. 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Biomim.</title><idno type="pISSN">1748-3182</idno><imprint><publisher>IOP Publishing</publisher><date type="published" when="2010"></date><biblScope unit="volume">5</biblScope><biblScope unit="issue">3</biblScope><biblScope unit="page" from="1">1</biblScope><biblScope unit="page" to="11">11</biblScope><biblScope unit="production">Printed in the UK</biblScope></imprint></monogr><idno type="istex">9ACE70A18185FE1AA829B2846A52A448DEC8EDA0</idno><idno type="DOI">10.1088/1748-3182/5/3/035006</idno><idno type="PII">S1748-3182(10)52493-1</idno><idno type="articleID">352493</idno><idno type="articleNumber">035006</idno></biblStruct></sourceDesc></fileDesc><profileDesc><creation><date>2010</date></creation><langUsage><language ident="en">en</language></langUsage><abstract><p>Behavioral and kinematic properties and capacities of wild migratory salmonid fishes swimming upstream and jumping up waterfalls generally have played only minor roles in the design and construction of passageways intended to help these fishes get past dams and other human-made obstacles blocking their movements. This paper reports the results of an experimental study of relevant behavioral and kinematic properties of adult kokanee salmon (Oncorhynchus nerka) jumping up waterfalls as they migrate upstream. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. Kokanee salmon spontaneously jump up waterfalls within a relatively narrow range of conditions, including low flow speeds, near vertical angles and pool depth to fall height ratios near 1.0. Preferred values for each parameter are, to some extent, dependent on other parameters. In contrast to previous misconceptions, jumping behavior is initiated by running S-start accelerations from beneath the boils formed in the plunge pools below waterfalls, as opposed to C-start standing jumps from the surface. S-starts are immediately followed by burst swimming to the point of takeoff at the surface. These results can contribute to an improved basis for developing designs of fish passageways that may ultimately make them more effective and efficient.</p></abstract><textClass><keywords scheme="keyword"><list><head>keywords</head><item><term>salmon</term></item><item><term>kokanee</term></item><item><term>jumping</term></item><item><term>waterfalls</term></item><item><term>behavior</term></item><item><term>kinematics</term></item><item><term>fish passageways</term></item><item><term>restoration ecology</term></item></list></keywords></textClass></profileDesc><revisionDesc><change when="2010">Published</change></revisionDesc></teiHeader></istex:fulltextTEI><json:item><extension>txt</extension><original>false</original><mimetype>text/plain</mimetype><uri>https://api.istex.fr/document/9ACE70A18185FE1AA829B2846A52A448DEC8EDA0/fulltext/txt</uri></json:item></fulltext><metadata><istex:metadataXml wicri:clean="corpus iop not found" wicri:toSee="no header"><istex:xmlDeclaration>version="1.0" encoding="ISO-8859-1"</istex:xmlDeclaration><istex:docType SYSTEM="http://ej.iop.org/dtd/iopv1_5_2.dtd" name="istex:docType"></istex:docType><istex:document><article artid="bb352493"><article-metadata><jnl-data jnlid="bb"><jnl-fullname>Bioinspiration & Biomimetics</jnl-fullname><jnl-abbreviation>Bioinsp. Biomim.</jnl-abbreviation><jnl-shortname>BB</jnl-shortname><jnl-issn>1748-3182</jnl-issn><jnl-coden>BBIICI</jnl-coden><jnl-imprint>IOP Publishing</jnl-imprint><jnl-web-address>stacks.iop.org/BB</jnl-web-address></jnl-data><volume-data><year-publication>2010</year-publication><volume-number>5</volume-number></volume-data><issue-data><issue-number>3</issue-number><coverdate>September 2010</coverdate></issue-data><article-data><article-type type="paper" sort="regular"></article-type><type-number type="paper" numbering="article" artnum="035006">035006</type-number><article-number>352493</article-number><first-page>1</first-page><last-page>11</last-page><length>11</length><pii>S1748-3182(10)52493-1</pii><doi>10.1088/1748-3182/5/3/035006</doi><copyright>2010 IOP Publishing Ltd</copyright><ccc>1748-3182/10/035006+11$30.00</ccc><printed>Printed in the UK</printed><features colour="none" mmedia="no" suppdata="no"></features></article-data></article-metadata><header><title-group><title>Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design</title><short-title>Salmon jumping</short-title><ej-title>Salmon jumping</ej-title></title-group><author-group><author address="bb352493ad1" email="bb352493ea1"><first-names>D V</first-names><second-name>Lauritzen</second-name></author><author address="bb352493ad2" email="bb352493ea2"><first-names>F S</first-names><second-name>Hertel</second-name></author><author address="bb352493ad1" email="bb352493ea3"><first-names>L K</first-names><second-name>Jordan</second-name></author><author address="bb352493ad1" email="bb352493ea4"><first-names>M S</first-names><second-name>Gordon</second-name></author></author-group><address-group><address id="bb352493ad1"><orgname>Department of Ecology and Evolutionary Biology, University of California</orgname>, Los Angeles, CA 90095-1606, <country>USA</country></address><address id="bb352493ad2"><orgname>Department of Biology, California State University—Northridge</orgname>, Northridge, CA 91330, <country>USA</country></address><e-address id="bb352493ea1"><email mailto="dlauritz@ccsf.edu">dlauritz@ccsf.edu</email></e-address><e-address id="bb352493ea2"><email mailto="fritz.hertel@csun.edu">fritz.hertel@csun.edu</email></e-address><e-address id="bb352493ea3"><email mailto="msgordon@ucla.edu">msgordon@ucla.edu</email></e-address><e-address id="bb352493ea4"><email mailto="ljordan@ucla.edu">ljordan@ucla.edu</email></e-address></address-group><history received="1 April 2010" accepted="29 July 2010" online="20 August 2010"></history><abstract-group><abstract><heading>Abstract</heading><p indent="no">Behavioral and kinematic properties and capacities of wild migratory salmonid fishes swimming upstream and jumping up waterfalls generally have played only minor roles in the design and construction of passageways intended to help these fishes get past dams and other human-made obstacles blocking their movements. This paper reports the results of an experimental study of relevant behavioral and kinematic properties of adult kokanee salmon (<italic>Oncorhynchus nerka</italic>) jumping up waterfalls as they migrate upstream. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. Kokanee salmon spontaneously jump up waterfalls within a relatively narrow range of conditions, including low flow speeds, near vertical angles and pool depth to fall height ratios near 1.0. Preferred values for each parameter are, to some extent, dependent on other parameters. In contrast to previous misconceptions, jumping behavior is initiated by running S-start accelerations from beneath the boils formed in the plunge pools below waterfalls, as opposed to C-start standing jumps from the surface. S-starts are immediately followed by burst swimming to the point of takeoff at the surface. These results can contribute to an improved basis for developing designs of fish passageways that may ultimately make them more effective and efficient.</p></abstract></abstract-group><classifications><keywords><keyword>salmon</keyword><keyword>kokanee</keyword><keyword>jumping</keyword><keyword>waterfalls</keyword><keyword>behavior</keyword><keyword>kinematics</keyword><keyword>fish passageways</keyword><keyword>restoration ecology</keyword></keywords></classifications></header><body numbering="bysection" refstyle="alphabetic"><sec-level1 id="bb352493s1" label="1"><heading>Introduction</heading><p indent="no">Most juvenile to adult salmonid fishes are spontaneous jumpers, often leaping high out of the water even when there appears to be no specific cause for this behavior. Adult wild salmonids often use this behavior as they try to pass waterfalls and other obstacles encountered during upstream spawning migrations. Humans also take the behavior into limited account in the designs of fish passageways intended to help migrating wild fish overcome otherwise impassable obstacles like weirs and dams (recent reviews and design specification documents include CDFO (<cite linkend="bb352493bib06" show="year">1980</cite>), Evans and Johnston (<cite linkend="bb352493bib11" show="year">1980</cite>), Beach (<cite linkend="bb352493bib01" show="year">1984</cite>), Clay (<cite linkend="bb352493bib08" show="year">1995</cite>), Flosi <italic>et al</italic> (<cite linkend="bb352493bib13" show="year">1998</cite>), Jungwirth <italic>et al</italic> (<cite linkend="bb352493bib15" show="year">1998</cite>), NMFS (<cite linkend="bb352493bib25" show="year">2008</cite>)).</p><p>Human-built fish passageways are, unfortunately, rarely adequately effective and efficient. This is the case for both downstream (juvenile) and upstream (adult) migrants. The passageways are more often seriously destructive to the fishes (Bowman <cite linkend="bb352493bib02" show="year">1996</cite>, Kareiva <italic>et al</italic> <cite linkend="bb352493bib16" show="year">2000</cite>, Ovidio and Philippart <cite linkend="bb352493bib27" show="year">2002</cite>, Holthe <italic>et al</italic> <cite linkend="bb352493bib14" show="year">2005</cite>, Caudill <italic>et al</italic> <cite linkend="bb352493bib07" show="year">2007</cite>, Ovidio <italic>et al</italic> <cite linkend="bb352493bib26" show="year">2007</cite>, Schilt <cite linkend="bb352493bib30" show="year">2007</cite>, Workman <cite linkend="bb352493bib41" show="year">2007</cite>, Lundqvist <italic>et al</italic> <cite linkend="bb352493bib20" show="year">2008</cite>, Thorstad <italic>et al</italic> <cite linkend="bb352493bib35" show="year">2008</cite>). They are usually well designed from engineering, economic and sometimes political perspectives, but most designs lack adequate consideration for fish behaviors, swimming biomechanics and kinematics, and sensory physiology (Clay <cite linkend="bb352493bib08" show="year">1995</cite>, NMFS <cite linkend="bb352493bib25" show="year">2008</cite>).</p><p>Several studies have explored aspects of fish (primarily salmonid) jumping behavior and associated swimming and jumping kinematics in the context of upstream movements through passageways (Denil <cite linkend="bb352493bib10" show="year">1937</cite>, McLeod and Nemenyi <cite linkend="bb352493bib22" show="year">1941</cite>, Collins and Elling <cite linkend="bb352493bib09" show="year">1960</cite>, Stuart <cite linkend="bb352493bib32" show="year">1962</cite>, Thompson <cite linkend="bb352493bib34" show="year">1970</cite>, Holthe <italic>et al</italic> <cite linkend="bb352493bib14" show="year">2005</cite>). Important limitations on most of this research are that the studies were done under laboratory conditions, often used small, non-migratory fish in small sized flumes, and did not include direct observations of how the fish jumped (Stuart <cite linkend="bb352493bib32" show="year">1962</cite>, Kondratieff and Myrick <cite linkend="bb352493bib17" show="year">2005</cite>, <cite linkend="bb352493bib18" show="year">2006</cite>, Brandt <italic>et al</italic> <cite linkend="bb352493bib03" show="year">2005</cite>). Holthe <italic>et al</italic> (<cite linkend="bb352493bib14" show="year">2005</cite>) used wild caught fishes under more realistic field conditions.</p><p>Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>) observed migrating wild sea run (anadromous) adult sockeye salmon (<italic>Oncorhynchus nerka</italic>) spontaneously leaping up natural waterfalls in Alaska. They showed that these fish jump in predictable ways and have specific preferences for jumping locations.</p><p>This paper reports the results of a follow-up experimental study of relevant behavioral and swimming kinematic properties of adult wild kokanee salmon belonging to an introduced California population as they jumped up artificial waterfalls during their upstream migration. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. The results can be useful in developing better designs for fish passageways. The study also raises the possibility of expanded roles, in carefully selected situations, for kokanee salmon in sustainable restoration of salmon runs in suitable habitats above existing dams.</p><p>Readers should keep in mind two important limitations on the work reported: (i) no work was done with juvenile fish migrating downstream and (ii) we did not control environmental temperatures. Seasonal variations in water temperatures have substantial effects on jumping abilities of different species (Holthe <italic>et al</italic> <cite linkend="bb352493bib14" show="year">2005</cite>) as well as more general effects on many other aspects of salmonid performance (Brett <cite linkend="bb352493bib04" show="year">1964</cite>, Myrick and Cech <cite linkend="bb352493bib24" show="year">2000</cite>). Significant evidence is accumulating that changes in water temperatures resulting from global climate change are likely to have important adverse effects on salmonid migrations (Farrell <cite linkend="bb352493bib12" show="year">2009</cite>).</p></sec-level1><sec-level1 id="bb352493s2" label="2"><heading>Methods</heading><sec-level2 id="bb352493s2-1" label="2.1"><heading>Animals</heading><p indent="no">We studied approximately 350 wild adult kokanee salmon (<italic>Oncorhychus nerka)</italic> (California Academy of Sciences Research Collection voucher specimen # CAS 85282) at Taylor Creek, Lake Tahoe Basin, California (38° 55′ 49″ N, 120° 3′ 16″ W) during their spawning migration in October 2000. The term kokanee refers to a large number of distinct sockeye salmon populations that are confined to fresh water and are not anadromous (they do not have a period of life in the sea). Fish in such populations are usually significantly smaller in size as adults than sea run fish. Other aspects of their life history are typical for the species (Burgner <cite linkend="bb352493bib05" show="year">1991</cite>).</p><p>Taylor Creek kokanee were chosen as study animals because of both their smaller adult size (making them easier to handle) and their accessibility during spawning migration. Average total length (TL) of fish studied was 29 cm (SD = 2, <italic>n</italic> = 21) and average mass 0.28 kg (SD = 0.05, <italic>n</italic> = 21). Adult sea run Alaska sockeye salmon studied previously averaged 59 cm TL and 2.8 kg mass (Lauritzen <italic>et al</italic> <cite linkend="bb352493bib19" show="year">2005</cite>).</p><p>Kokanee spawning runs in Taylor Creek (3.3 km long) are large, up to totals of near 80 000 individuals (estimate per California Department of Fish and Game; CDFG). This results in high densities of fish that may be easily obtained for study. Unlike most of California's Pacific salmon populations, Tahoe kokanee are not listed as either threatened or endangered. The Lake Tahoe population is hatchery based. It is maintained by CDFG, which made the original introductions in 1943–1944. The present population has multiple geographic origins, as some new fish are introduced almost annually to maintain genetic diversity.</p><p>All collections were authorized with a CDFG Scientific Collecting permit. All handling of study animals was done in accordance with University of California Los Angeles Animal Research Committee guidelines (ARC protocol #98-169-03).</p></sec-level2><sec-level2 id="bb352493s2-2" label="2.2"><heading>Experimental apparatus: <italic>artificial waterfall generator</italic></heading><p indent="no">All experiments were done using a custom-designed and-built movable flume and artificial waterfall generator (AWG; figure <figref linkend="bb352493fig01">1</figref>). The AWG was set up in the streambed of Taylor Creek. All experiments were done using water directly from the Creek, under ambient environmental conditions. The AWG was adjustable in multiple ways: volumes and speeds of water flows; heights of artificial falls; angles of channels and falls; and dimensions of the observation pool.
<figure id="bb352493fig01" width="page"><graphic><graphic-file version="print" format="EPS" filename="images/bb352493fig01.eps" width="26pc"></graphic-file><graphic-file version="ej" format="JPEG" filename="images/bb352493fig01.jpg"></graphic-file></graphic><caption id="bb352493fc01" label="Figure 1"><p indent="no">A schematic diagram of the AWG apparatus with the three systems labeled. Gray shading represents water. A: drain, B: pool outflow spout, C: false bottom, D: viewing pool, E: outflow spout, F: water column chute, G: outflow spout, H: water velocity reducing column, I: volume flux control valve, J: discharge hose, K: water pump, L: suction hose, M: fish screen, N: water column support, O: water channel support, P: water channel, Q: ramp at 90°.</p></caption></figure></p><sec-level3 id="bb352493s2-2-1" label="2.2.1"><heading><italic>Artificial waterfall generator</italic> design and construction</heading><p indent="no">The AWG had three parts: a water delivery system (WDS), a water transfer system (WTS) and an observation pool (OP). Using pumps, the WDS drew water from Taylor Creek. Water was guided to the adjustable artificial waterfall by the WTS. Water poured over the fall and into the OP, where fish were introduced and experimental trials initiated. The downstream end of the OP allowed for water to over-spill back into the Creek.</p><p>Refer to letters in figure <figref linkend="bb352493fig01">1</figref> to follow this description. Primary materials used in the AWG were Varathane (Flecto) sealed wood, with steel supports and metal fasteners.</p><p><italic>WDS</italic>. The WDS was driven by two 8 hp gasoline powered water pumps (K; Honda Power Equipment model WT30X). These pumps drew water through non-collapsible suction hoses (L, M; 7.6 cm diameter) fitted with fish/debris screens on the intake ends. Collapsible discharge hoses (J; 7.6 cm diameter) connected the pumps to the water column that led to the WTS.</p><p>The most prominent feature of the WDS was the water velocity reducing column, consisting of two polyethylene (PE) open-head 208 liter (l) drums (H) fastened to each other at the open ends. Intake fittings for the discharge hoses were mounted on this column midway up the lower drum. Internal intake directional nozzles were fastened to the intake fittings and oriented downward at 45° angles. These nozzles assisted in reducing the high velocities of water jetting from the discharge hoses.</p><p>The lower drum was also fitted with a 7.6 cm diameter polyvinyl chloride (PVC) ball valve (I) that acted as the volume control. An outflow spout made of stainless steel (G) was located at the top of the upper drum. This spout directed water flow to the water column chute (F; 114 l PE closed-head drum). The height of the chute was adjustable so it could match the height of the water channel. Flow out of the chute was directed with a second stainless steel outflow spout (E).</p><p>The entire water column system was supported by a wooden structure with steel brackets, tubing and threaded fasteners (N). The support structure was designed to support the water column system under full water loads with a seismic safety factor of 1 g.</p><p><italic>WTS</italic>. The WTS contained the water channel (P; 135 × 30 × 14 cm) that caught water leaving the water column chute and delivered it to the OP via the waterfall. The channel was open at the downstream end to create a waterfall and closed at the upstream end to prevent back flow. The channel was supported (O) by four vertical steel tubes attached to two horizontal wooden beams that were oriented cross-wise to the axis of the apparatus. Steel pins connected the tubes to these beams at a range of positions along the lengths of the tubes; this allowed for adjustment of waterfall height. The transverse beams were movable; they could slide forward or back on top of two other horizontal beams (oriented parallel to the axis of the apparatus) that were mounted between the water column support and the OP. This up and downstream sliding of the channel allowed for variability in the gradient of the falls.</p><p>Two different ramps could be attached to the downstream end of the water channel. The vertical ramp (Q) consisted of a hanging wooden sheet (61 × 30 cm) attached by a hinge to the water channel. This ramp acted as a backdrop for vertical waterfall gradients. The incline ramp was a channel (61 × 30 cm) with sidewalls (14 cm high) that was open at both ends. This ramp could be attached to both the water channel and the OP by hinges; it was used to produce fall gradients at angles below vertical.</p><p><italic>OP</italic>. The waterfall at the end of the WTS drained into the OP. The OP was a large wooden tank (224 × 71 × 117 cm) reinforced around the walls and across the top with a steel pipe. It was fastened together with carriage bolts, lag bolts and square-drive deck screws. A weighted wooden sheet (C) acted as a movable false bottom to the pool. The sheet could be anchored at various depths in the pool using four nylon lines tied off to four cleats on the outside top corners of the tank.</p><p>The upstream halves of the sidewalls of the OP (D) were made of clear acrylic panels (1.9 cm thick) for viewing fish underwater. The top downstream edge of the pool housed the pool outflow spout of the apparatus (B). This spout had an adjustable aperture that made it possible to maintain pool height over a variety of flow rates. The bottom of the downstream wall also contained a 7.6 cm PVC drain plug (A) to allow for rapid draining of the pool.</p></sec-level3><sec-level3 id="bb352493s2-2-2" label="2.2.2"><heading>Flow volumes, speeds and circulation patterns in the observation pool</heading><p indent="no">Volumes of water flowing over the fall and into the OP varied with changes in fall height, fall gradient, pool depth and flow speed. Water flows separated from the WTS at the crest of the fall at all flow rates with a vertical ramp angle. Separations did not occur at ramp angles significantly less than vertical.</p><p>Fall gradients of 0° produced surface currents in the OP oriented to the outflow spout while simultaneously producing recirculating flows beneath this current (figure <figref linkend="bb352493fig02" override="yes">2(<italic>A</italic>)</figref>). Vertical fall gradients (90°) produced steep downward-directed currents that were redirected back toward the surface by the false bottom. This formed a boil of water at the surface of the pool immediately downstream of the falls. Water then flowed along the pool surface toward the pool outflow (figure <figref linkend="bb352493fig02" override="yes">2(<italic>B</italic>)</figref>). Intermediate fall gradients produced combinations of these two flow schemes. Deeper pool depths required higher flow rates and higher fall heights to produce surface boils in the OP.
<figure id="bb352493fig02"><graphic><graphic-file version="print" format="EPS" filename="images/bb352493fig02.eps" width="18pc"></graphic-file><graphic-file version="ej" format="JPEG" filename="images/bb352493fig02.jpg"></graphic-file></graphic><caption id="bb352493fc02" label="Figure 2"><p indent="no">Flow patterns in the observation pool are shown as black arrows with a ramp angle of A 0° and B 90°. Gray shading represents water.</p></caption></figure></p><p>Both volumes and speeds of water flowing through the system under different conditions were estimated by calculation. The calculated velocity of the water at the instant it poured out of the upstream outflow spout of the WDS assumed that water poured from the spout following a parabolic trajectory (Mulligan <cite linkend="bb352493bib23" show="year">1991</cite>); no correction was made for possible air resistance. We also assumed that the velocity was solely composed of a horizontal component (the vertical velocity component at this moment was zero). The horizontal velocity component equaled the known distance traveled (acquired with a measuring tape) divided by the time taken to travel that distance. The time was calculated with the parabolic function describing an object (a volume of water) free falling for the same duration that the water was displaced horizontally. This function simplifies to <italic>t</italic> = (2 h g<sup>−1</sup>)<sup>0.5</sup> where <italic>t</italic> is the time traveled, <italic>h</italic> is the height from which the object drops and <italic>g</italic> is the gravitational constant (980 cm s<sup>−2</sup>). Volume flux was calculated by multiplying the velocity by the cross-sectional area of water coming out of the upstream outflow spout.</p></sec-level3><sec-level3 id="bb352493s2-2-3" label="2.2.3"><heading>Adjustability</heading><p indent="no">The AWG was capable of providing a wide range of experimental conditions by producing a variety of waterfalls. Four adjustable parameters were incorporated into the design. The height of the waterfall was adjustable from 0 to 72 cm at increments of 12 cm by raising or lowering the water channel. The depth of the false bottom in the OP controlled pool depth below the fall. Pool depths ranged from 8 to 114 cm with a continuous range of intermediate depths. The gradient of the fall was set by changing the angle of the incline ramp that directed water down the fall and into the pool. Ramp angles could be set at nine different values (0, 8, 18, 28, 39, 51, 63, 76 and 90°). The rate of water flow over the fall was controlled with the throttle speed on the pumps that drove the apparatus. With one pump running, the water velocity pouring out of the WDS ranged from 0 to 1.1 m s<sup>−1</sup> and the volume flux through the system ranged from 0 to 13 L s<sup>−1</sup>.</p></sec-level3></sec-level2><sec-level2 id="bb352493s2-3" label="2.3"><heading>Experimental trials</heading><p indent="no">Trials consisted of both visually observing and videotaping spontaneous jumps of fish up the artificial waterfall. Variables included flow rates, pool depths, fall heights and fall gradients. The protocol followed in each trial was as follows.</p><p>Ten fish were placed together in the OP. After 5 min acclimation periods, observations were made for a minimum of 15 min. If no fish attempted to jump up the fall during that time the trial was ended. If one or more attempts were made during the first 15 min, observations continued for 10 min after the first attempt was noted. Attempts were defined as either an individual fish jumped clear of the water or the pectoral fins of a fish entered the incline ramp.</p><p>Experiments were run under a variety of configurations to determine which sets of conditions elicited jumping behavior. Trials were run at three different flow rates (3.8 L s<sup>−1</sup>, 93 cm s<sup>−1</sup>; 11.5 L s<sup>−1</sup>, 109 cm s<sup>−1</sup>; 12.8 L s<sup>−1</sup>, 114 cm s<sup>−1</sup>; table <tabref linkend="bb352493tab01">1</tabref>). Flow rates used corresponded to one pump running at idle, half throttle and full throttle. Seven different pool depths were used (8, 15, 23, 30, 38, 46 and 66 cm). Depths were set using a measuring tape from the surface of the pool water to the false bottom suspended in the pool. Fall heights were set at 0, 13, 25 and 36 cm. Heights were measured with a measuring tape from the surface of the pool water to the crest of the falls. Fall gradients were set at 0, 18, 38, 64 and 90° from the horizontal. Gradient angles were measured using digital images taken from the video database and ImageJ version 1.24t software (NIH: <webref url="http://rsb.info.nih.gov/ij">http://rsb.info.nih.gov/ij</webref>). For each set of conditions, the number of attempts was noted. When multiple trials with identical conditions were run, their average results were used in calculating preferences. The set of conditions that produced the highest rate of attempts is considered the preferred set of conditions among those compared.
<table id="bb352493tab01" frame="topbot"><caption id="bb352493tc01" label="Table 1"><p indent="no">Fish advancing up the falls at three pump speeds. Numbers of jumps are from trials at four pool depths (8, 15, 30 and 66 cm) and three fall heights (13, 25 and 36 cm) for totals of 12 trials at each pump speed. Water velocities scaled to body lengths s<sup>−1</sup>(BL s<sup>−1</sup>) based on 29 cm mean salmon total lengths.</p></caption><tgroup cols="4"><colspec colnum="1" colname="col1" align="left"></colspec><colspec colnum="2" colname="col2" align="left"></colspec><colspec colnum="3" colname="col3" align="left"></colspec><colspec colnum="4" colname="col4" align="left"></colspec><thead><row><entry>Pump</entry><entry>Volume</entry><entry>Water velocity</entry><entry>Number</entry></row><row><entry>speed</entry><entry>flux (L s<sup>−1</sup>)</entry><entry>(cm s<sup>−1</sup>; BL s<sup>−1</sup>)</entry><entry>of jumps</entry></row></thead><tbody><row><entry>Idle</entry><entry> 3.8</entry><entry> 93; 3.2</entry><entry>32</entry></row><row><entry>Half</entry><entry>11.5</entry><entry>109; 3.8</entry><entry> 1</entry></row><row><entry>Full</entry><entry>12.8</entry><entry>114; 3.9</entry><entry> 0</entry></row></tbody></tgroup></table></p></sec-level2><sec-level2 id="bb352493s2-4" label="2.4"><heading>Video recording and calibration</heading><p indent="no">The behavioral and kinematic database consisted of 7 h of digital video recordings. All trials for preference experiments and biomechanical analysis were recorded at standard speed (30 fps) with a DCR-PC100 camcorder (Sony Corporation, Tokyo, Japan). All trials for biomechanical analyses were recorded simultaneously at high speed (1000 fps) with a Motion Scope 1000S digital video camera (Redlake MSAD, Inc.). Videos were taken perpendicular to the flow of the water in the apparatus. Both cameras were mounted on the same tripod at a distance of 490 cm from the midline of the OP.</p><p>Twenty-five jumps recorded with high speed video and 38 with low speed video were used for kinematic analyses. Jump trajectories that resulted in a displacement of more than 15 cm in directions perpendicular to the flow of water were not analyzed in order to reduce effects of perspective on analyses and limit the error in calculated linear dimensions. Calibration of the hardware used in capturing video and transferring it to a personal computer is described in Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>). Individual fish positions throughout jumps were obtained using the ImageJ software. Mathematica version 3.0 software (Wolfram Research) was used to convert dimensions measured in the video frames from pixels to centimeters (using objects of known lengths visible in the frames of each jump) and to calculate jumping parameter values. For calibration, we used two objects at 1000 fps and three objects at 30 fps. Accuracy of the calibration method was checked against an additional known distance visible in recordings at both frame rates and was not significantly different (<italic>p</italic> > 0.05 for both tests, Student's <italic>t</italic>-distribution).</p></sec-level2><sec-level2 id="bb352493s2-5" label="2.5"><heading>Kinematic jumping parameters</heading><p indent="no">All data collected for kinematic study are based on jumps that took place with a pool depth of 30 cm, fall height of 36 cm, falls gradient of 90°, water volume flux of 3.8 L s<sup>−1</sup> and water velocity of 93 cm s<sup>−1</sup>. These parameters provided adequate views of both air and water in addition to a relatively high jumping rate (figure <figref linkend="bb352493fig03">3</figref>). Up to ten fish were in the OP during kinematic trials.
<figure id="bb352493fig03"><graphic><graphic-file version="print" format="EPS" filename="images/bb352493fig03.eps" width="20.5pc"></graphic-file><graphic-file version="ej" format="JPEG" filename="images/bb352493fig03.jpg"></graphic-file></graphic><caption id="bb352493fc03" label="Figure 3"><p indent="no">A kokanee salmon jumping from the OP to the crest of the waterfall. The video frame shows simultaneous movement of fish in water and air.</p></caption></figure></p><p>Takeoff velocities and angles were calculated from high speed video of jumping events. These parameter values are based on parabolic functions describing ballistic trajectories (Mulligan <cite linkend="bb352493bib23" show="year">1991</cite>) while neglecting air resistance (Lauritzen <italic>et al</italic> <cite linkend="bb352493bib19" show="year">2005</cite>) and were calculated using the same methods used by Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>). The remaining kinematic parameters were calculated using standard speed video. Pre-acceleration velocities were calculated by dividing the distance between the point of maximum depth attained during a jump approach and the start of a rapid acceleration toward the pool surface by the time required to travel this distance. The horizontal distances traveled between takeoff and the falls crest were also measured. All comparisons of means were made using the Student's t-distribution with a level of significance at <italic>p</italic> < 0.05.</p></sec-level2></sec-level1><sec-level1 id="bb352493s3" label="3"><heading>Results</heading><sec-level2 id="bb352493s3-1" label="3.1"><heading>Range of observed behaviors</heading><p indent="no">Fish in the OP behaved in many different ways. There were five different types of responses to water flows: (1) flow avoidance; (2) lack of visible response to flow; (3) station holding; (4) swimming up the ramp and (5) jumping up the fall.</p><p>Fish responding in any of the first three ways did not advance up the fall. Many fish avoided high flows pouring into the pool by quickly swimming to the lower upstream corners of the pool where they usually remained throughout their trials (figure <figref linkend="bb352493fig04">4</figref>A). During trials at low flow rates fish typically distributed themselves throughout the pool, apparently ignoring the flow. This happened most often in trials using deeper pools. Stronger flows and/or shallower pool depths forced fish to display more distinct responses to flow. Fish often demonstrated classic rheotactic responses by orienting into the flow and maintaining position (station holding). This was the most commonly observed behavior in this study (figure <figref linkend="bb352493fig04">4</figref>B).
<figure id="bb352493fig04"><graphic><graphic-file version="print" format="EPS" filename="images/bb352493fig04.eps" width="20.5pc"></graphic-file><graphic-file version="ej" format="JPEG" filename="images/bb352493fig04.jpg"></graphic-file></graphic><caption id="bb352493fc04" label="Figure 4"><p indent="no">Diagram of four of the fish behaviors seen in the OP. Flow avoidance during high flow rates (A); station holding in the current (B); swimming up the shallow angle diagonal ramp (C) and jumping up the high angle ramp or vertical fall (D). Black dots represent trajectories of fish C and D. The spacing between consecutive dots represents relative velocities at those positions within the trajectories.</p></caption></figure></p><p>Trials with shallow ramp angles frequently induced fish to swim up on the ramp. Swimming attempts were always initiated from the base of the ramp with little or no burst swimming approach. Approaches to the base of the ramp were usually leisurely and started beneath the flow coming off the ramp (figure <figref linkend="bb352493fig04">4</figref>C).</p><p>Steeper ramp angles, including vertical angles, resulted in fish jumping up the ramp. These jumps involved running starts. Jumps were initiated downstream from the boil produced by water pouring over the falls. Fish slowly swam up to the base of the boil near the bottom of the pool and rotated their bodies to orient their heads so that they faced the pool surface. Fish then made S-starts, first arching their bodies, then rapidly beat their tails and accelerated through the boil to takeoff. Upon takeoff, fish traveled through the air toward the crest of the falls while continuing to beat their caudal fins (figure <figref linkend="bb352493fig04" override="yes">4(<italic>D</italic>)</figref>).</p><p>Very few jumps reached the crest of the falls (or higher) and even fewer were successful in topping the falls. Successful jumping fish were those that were able to momentarily maintain position or advance up the ramp upon landing and not be immediately pushed back down into the pool. The only jumps that were successful were those that landed above the crest of the falls. Of the 193 jumps analyzed, 35 were successful, yielding an 18% success rate.</p></sec-level2><sec-level2 id="bb352493s3-2" label="3.2"><heading>Jump-inducing conditions</heading><p indent="no">The rate of water flowing through the apparatus had a strong effect on the jumping tendencies of fish. Of the 12 trials run at each flow rate, fish were much more likely to jump up the vertical ramp at the lowest pump speed (32 jumps observed) than at the middle (1 jump observed) or highest (no jumps observed) pump speeds (table <tabref linkend="bb352493tab01">1</tabref>). When analyzed separately, ranges of pool depths and fall heights did not have statistically significant effects on fish jumping. The mean numbers of jumps observed (taken from three different fall heights: 13, 25 and 36 cm) for each depth tested (8, 15, 23, 30 and 66 cm) were not significantly different from each other (<italic>p</italic> > 0.05; figure <figref linkend="bb352493fig05" override="yes">5(<italic>A</italic>)</figref>). Similarly, mean numbers of jumps observed (taken from five different pool depths: 8, 15, 23, 30 and 66 cm) for each fall height tested (13, 25 and 36 cm) were not significantly different from each other (<italic>p</italic> > 0.05; figure <figref linkend="bb352493fig05" override="yes">5(<italic>B</italic>)</figref>). An effect of the pool depth and fall height on jumping behavior is seen when considering both characteristics simultaneously. The preferred pool depth for jumping slightly increases with an increase in the fall height (figure <figref linkend="bb352493fig05" override="yes">5(<italic>C</italic>)</figref>), but the preferred ratio of the pool depth to fall height (<italic>D</italic>/<italic>H</italic>) decreases with an increase in the fall height (fall height = 13 cm, <italic>D</italic>/<italic>H</italic> = 1.2; fall height = 25 cm, <italic>D</italic>/<italic>H</italic> = 0.9; fall height = 36 cm, <italic>D</italic>/<italic>H</italic> = 0.6).
<figure id="bb352493fig05" width="page"><graphic><graphic-file version="print" format="EPS" filename="images/bb352493fig05.eps" width="42pc"></graphic-file><graphic-file version="ej" format="JPEG" filename="images/bb352493fig05.jpg"></graphic-file></graphic><caption id="bb352493fc05" label="Figure 5"><p indent="no">Physical conditions influencing the number of jumps by fish. (<italic>A</italic>) Jump variation with the pool depth. Circles represent the means from three trials at varying fall heights. Error bars are one standard deviation from the means. (<italic>B</italic>) Jump variation with fall heights. Circles represent the means from five trials with varying pool depths. Error bars are one standard deviation from the mean. (<italic>C</italic>) Numbers of jumps observed at seven pool depths for three fall heights. Hollow circles represent a fall height of 13 cm, solid circles represent a fall height of 25 cm, and solid squares represent a fall height of 36 cm. Lines do not represent continuity between data points but instead are aids in visualizing data points for each fall height. (<italic>D</italic>) Proportion of jump attempts plotted against ramp angles. A best fit sigmoidal function shows an inflection point near 40°. We assume that all attempts at a ramp angle of 0° are swimming and all attempts at 90° are jumps. These assumptions were validated by observed fish behavior. The shaded area represents jumping attempts and the unshaded area represents swimming attempts.</p></caption></figure></p><p>The proportions of attempts to get up the ramp that were made by jumping, instead of swimming, were calculated for five ramp angles ranging from 0 to 90° (table <tabref linkend="bb352493tab02">2</tabref>). The ramp angles versus the jump proportions form a sigmoidal relationship with an inflection point near 40° (figure <figref linkend="bb352493fig05" override="yes">5(<italic>D</italic>)</figref>).
<table id="bb352493tab02" frame="topbot"><caption id="bb352493tc02" label="Table 2"><p indent="no">Numbers of attempts made by fish to move up the ramp and the fraction of those attempts that were jumps at each of five ramp angles.</p></caption><tgroup cols="3"><colspec colnum="1" colname="col1" align="left"></colspec><colspec colnum="2" colname="col2" align="left"></colspec><colspec colnum="3" colname="col3" align="left"></colspec><thead><row><entry>Ramp</entry><entry>Numbers</entry><entry>Jump</entry></row><row><entry>angles (°)</entry><entry>of attempts</entry><entry>fractions</entry></row></thead><tbody><row><entry> 0</entry><entry>11</entry><entry>0.00</entry></row><row><entry>18</entry><entry>18</entry><entry>0.11</entry></row><row><entry>38</entry><entry> 7</entry><entry>0.43</entry></row><row><entry>64</entry><entry> 9</entry><entry>1.00</entry></row><row><entry>90</entry><entry>16</entry><entry>1.00</entry></row></tbody></tgroup></table></p><p>To summarize, preferred jumping conditions are at low flow speeds with vertical ramp angles and pool depth to height ratios near 1.0.</p></sec-level2><sec-level2 id="bb352493s3-3" label="3.3"><heading>Kinematic jumping parameters</heading><p indent="no">Takeoff velocities and angles were calculated from 25 jumps captured with high speed video. The other parameters, including depths, distances and pre-acceleration velocities, were calculated from the same 25 jumps plus 14 additional jumps. Jumps observed in this study were powered by rapid starts near the bottom of the pool beneath the boil in a similar manner to the jumps observed in wild Alaska salmon (Lauritzen <italic>et al</italic> <cite linkend="bb352493bib19" show="year">2005</cite>). Fish slowly approached the bottom of the pool beneath the boil where rapid accelerations were initiated. Mean pre-acceleration velocities were 21 cm s<sup>−1</sup>, which increased to 229 cm s<sup>−1</sup> at takeoff with the body at an angle of 60° (table <tabref linkend="bb352493tab03">3</tabref>). These accelerations were produced by rapidly bending their bodies and were followed by burst swimming to the takeoff points at the surface of the boil.
<table id="bb352493tab03" frame="topbot"><caption id="bb352493tc03" label="Table 3"><p indent="no">Kinematic jumping parameters calculated from standard speed (SS; <italic>N</italic> = 38 fish) and high speed (HS; <italic>N</italic> = 25 fish) video recordings. All data collected from trials with water flow rates of 3.8 L s<sup>−1</sup>, pool depths of 30 cm and ramp angles of 90°.</p></caption><tgroup cols="3"><colspec colnum="1" colname="col1" align="left"></colspec><colspec colnum="2" colname="col2" align="left"></colspec><colspec colnum="3" colname="col3" align="left"></colspec><spanspec namest="col1" nameend="col3" spanname="1to3" align="center"></spanspec><thead><row><entry></entry><entry></entry><entry>Standard</entry></row><row><entry>Parameter (SS videos)</entry><entry>Mean</entry><entry>deviation</entry></row></thead><tbody><row><entry>Maximum depth (cm)</entry><entry> 21</entry><entry> 4</entry></row><row><entry>Horizontal distance traveled (cm)</entry><entry> 24</entry><entry>12</entry></row><row><entry>Pre-acceleration speed (cm s<sup>−1</sup>)</entry><entry> 21</entry><entry> 8</entry></row><row><entry>Depth at the start of acceleration (cm)</entry><entry> 13</entry><entry> 5</entry></row><row><entry>Parameter (HS videos)</entry><entry></entry><entry></entry></row><row><entry spanname="1to3"></entry></row><row><entry>Horizontal velocity (cm s<sup>−1</sup>)</entry><entry>114</entry><entry>46</entry></row><row><entry>Vertical takeoff velocity (cm s<sup>−1</sup>)</entry><entry>195</entry><entry>54</entry></row><row><entry>Total takeoff velocity (cm s<sup>−1</sup>)</entry><entry>229</entry><entry>59</entry></row><row><entry>Takeoff angle (°)</entry><entry> 60</entry><entry>11</entry></row></tbody></tgroup></table></p></sec-level2></sec-level1><sec-level1 id="bb352493s4" label="4"><heading>Discussion</heading><sec-level2 id="bb352493s4-1" label="4.1"><heading>Implications</heading><p indent="no">These results have several different implications:
<ordered-list id="bb352493ol1" type="alpha" pattern="2"><list-item id="bb352493ol1.1" marker="(a)"><p indent="no">The clear preferences migratory kokanee salmon demonstrate for relatively narrow ranges of water flow conditions, waterfall heights and waterfall angles provide important parameters that should be taken into account in future designs of passageways for salmonids, or in programs of retrofitting existing passageways. Explicit provision should be made for their use of running S-starts, rather than standing C-starts.</p></list-item><list-item id="bb352493ol1.2" marker="(b)"><p indent="no">Further studies similar to this one would be valuable. They could provide data relating to additional important factors, such as adjustments in scaling of the parameters that would make them directly applicable to the much larger sea run salmonids; effects of varying water temperatures and differences in the parameters that undoubtedly exist between salmonids and other important migratory species, such as clupeid fishes (e.g. shad and alewives) and sturgeon.</p></list-item><list-item id="bb352493ol1.3" marker="(c)"><p indent="no">The kokanee salmon in the Lake Tahoe–Taylor Creek population have encountered no circumstances requiring jumping for at least the past 65+ years since they were introduced into the region. The salmon found in Taylor Creek today are mostly descendants of the original fish (Reiner, CDFG, personal communication). The short length of Taylor Creek accessible to them is almost level, and their upstream migration is stopped at its upper end by an impassable dam that has a poorly designed non-functional fish passageway that the fish do not use. The fact that they remain active jumpers despite this multi-generational lack of reinforcement of the behavior supports the conventional wisdom that the suite of biomechanical, sensory and motor capacities involved is evolutionarily derived and genetically conserved in their nervous systems. Our results show that, when presented with the proper flow conditions, these fish will readily jump up a waterfall even though this population of fish has not used this behavior for more than 25 generations.</p></list-item></ordered-list></p></sec-level2><sec-level2 id="bb352493s4-2" label="4.2"><heading>Jumping preferences</heading><p indent="no">Kokanee salmon exhibited several different responses to the variety of waterfall conditions presented. There was, however, only a narrow range of conditions that prompted them to jump. Very slow water velocities apparently did not provide a strong enough rheotactic cue for fish to jump. The importance of this minimal flow has been recognized for many years and is referred to as attraction water (Clay <cite linkend="bb352493bib08" show="year">1995</cite>). The flow rate ‘window’ within which jumping occurred was narrow and frequencies decreased rapidly with relatively small increases in water flow rates. Using water velocity increments of less than 1 BL s<sup>−1</sup>, all jumping activity ceased at velocities between 3 and 4 BL s<sup>−1</sup>.</p><p>The stoppage of jumping activity at flow rates above 3–4 BL s<sup>−1</sup> may possibly have been at least partly an artifact of the relatively short durations of our observation periods. A contributing factor could have been the larger energy demands associated with maintaining position (station holding) in higher flows. Some fish may have been somewhat fatigued by their efforts to maintain station. The corners of the OP provided easily accessible resting areas with low flows. Those corners were used by significant numbers of fish. The short durations of our observation periods may not have been long enough for them to fully recover from this fatigue, so they made no effort to jump. It is possible that longer recovery times could have resulted in jumping at higher flow rates. In addition, some fish may have been sufficiently stressed from recent handling while being placed into the OP that they were less active and responsive than they might have been.</p><p>Effects on jumping behavior of variable pool depths and fall heights are only apparent when one of these parameters is held constant. For each fall height, there was a range of pool depths where jumping behavior was observed (figure <figref linkend="bb352493fig05" override="yes">5(A)</figref>). Very few jumps were observed at the shallowest pool depth, 8 cm, and no jumps occurred at depths greater than 38 cm. Pool depths of 8 cm and less were too shallow for fish to generate adequate takeoff velocities oriented toward the crest of the fall. Pool depths greater than 38 cm appeared to allow fish to avoid the current below the fall.</p><p>Jumping approaches originated from downstream and were initiated by swimming toward the bottom of the boil below the fall. When the floor of the pool was considerably lower than the bottom of the boil, greater than a depth of 38 cm, fish would continue to swim downward as they reached the boil. Leaving the current generated by the fall resulted in aborted jumps. Deeper pools allowed fish to remain passively in the lower portion of the pool without being influenced by the current flowing through the system.</p><p>The preferred pool depth/fall height ratios for this study were 1.2, 0.9 and 0.6 for fall heights of 13, 25 and 36 cm, respectively. These ratios are all lower than the preferred ratio suggested by Stuart (<cite linkend="bb352493bib32" show="year">1962</cite>), but they overlap with the ratios observed in sea run salmon by Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>; ratios of 0.7 and 1.5). If a specific ratio does exist where the maximum number of jumps is induced in salmonids, that ratio is probably less than Stuart's proposed 1.25. Other species that have been tested also showed lower pool depth/fall height ratios: 1.1 for 26 cm FL (fork length) brown trout and 0.4 for both 29.5 cm FL brown trout and 27 cm FL grayling (Ovidio <italic>et al</italic> <cite linkend="bb352493bib26" show="year">2007</cite>). The average ratio we observed, 1.0, may be a more accurate value. This ratio lies well within the range of pool depths that induced jumping at all three fall heights in this study. However, the variability of these ratios suggests that other factors may also be involved in producing preferred jumping conditions.</p><p>The fall gradient is probably one of these factors. As the gradient of the fall increased from horizontal to vertical, attempts to reach the fall crest switched from 100% swimming to 100% jumping. The work against gravity required to swim up to the crest increased as the ramp angles increased. At the ramp angle where the work required to swim was equal to the work required to jump 50% of the attempts to get up the fall were by swimming and 50% were by jumping. This angle is noted in figure <figref linkend="bb352493fig05" override="yes">5(D)</figref> as the inflection point of the sigmoidal curve fitted through the data. Due to the drastically different approaches used by fish for these two behaviors, passageway pool designs must take into account both anticipated modes of travel. Jumping attempts require deeper and longer approaches, consequently deeper and longer pools, than do swimming attempts. Although jumping behavior requires larger pools, fish passageways designed for jumping fish (vertical falls) take up considerably less horizontal area for a given elevation gain.</p></sec-level2><sec-level2 id="bb352493s4-3" label="4.3"><heading>Jump kinematics</heading><p indent="no">Our results differ from previous understanding of how salmonids jump up falls. Stuart (<cite linkend="bb352493bib32" show="year">1962</cite>) states that salmon slowly rise to the pool surface and then use a C-start (single large arc in the body) to accelerate to takeoff velocity. The body bending observed in our kokanee produced sinusoidal waves in the fish, which more closely resembled an S-start type of rapid acceleration (Weihs <cite linkend="bb352493bib38" show="year">1972</cite>, <cite linkend="bb352493bib39" show="year">1973</cite>, Webb <cite linkend="bb352493bib36" show="year">1975</cite>, Webb and Skadsen <cite linkend="bb352493bib37" show="year">1980</cite>). The fish in this study left the water with similar body attitudes and takeoff angles as the sea run fish observed by Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>). Fish from both studies exited the pools with their bodies straight and their caudal fins beating, with a combined mean takeoff angle of 58° (no significant difference between the angles in these studies; <italic>p</italic> = 0.12). The similarities in how fish exited the pools in both studies suggest that the larger sea run salmon also used the same technique to reach takeoff velocities as the kokanee salmon in this study. By initiating the acceleration at the bottom of the pool followed by burst swimming, fish are able to reach higher takeoff velocities than if they just elicit a C-start-type startle response at the surface.</p><p>Additionally, S-starts are better suited for accelerations toward a specific target such as the crest of a waterfall. C-starts produce a large moment perpendicular to the original axis of the fish body resulting in a large lateral movement. S-start kinematics do not involve these large diversions and allow fish to accelerate in the original direction (Weihs <cite linkend="bb352493bib38" show="year">1972</cite>, <cite linkend="bb352493bib39" show="year">1973</cite>, Webb <cite linkend="bb352493bib36" show="year">1975</cite>, Webb and Skadsen <cite linkend="bb352493bib37" show="year">1980</cite>).</p><p>The mathematical model proposed by Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>) indicates that the takeoff velocity of fish jumping up falls should be primarily dependent on the height of the falls. Therefore, one would expect salmon of different populations to have similar takeoff velocities while jumping up falls of equal heights, when those heights are scaled to the sizes of the fish. The mean takeoff velocity of kokanee salmon jumping in the AWG (fall height = 1.2 BL) is nearly the same as the mean velocity of sockeye salmon jumping at Brooks Falls, Alaska (fall height = 1.4 BL). The difference in takeoff velocities between these fish was 0.7 BL s<sup>−1</sup> (<italic>p</italic> = 0.05).</p><p>The borderline <italic>p</italic> value for this comparison may be explained by a vectorial analysis of the directional (horizontal and vertical) components of the takeoff velocities (table <tabref linkend="bb352493tab03">3</tabref>). There is no significant difference between the vertical velocity components (<italic>p</italic> = 0.15), but there is a significant difference between the horizontal components (<italic>p</italic> < 0.05). The higher horizontal velocity component for the fish at Brooks Falls is a result of those fish jumping from farther downstream from the falls than the fish in the AWG. The discrepancy between horizontal takeoff distances from the falls may be due to differences in hydrodynamic signals perceived by the fish at different locations or due to evolutionary differences between the fish. The relatively low successful jumping rates observed in this study (18%) are similar to the rates reported by Lauritzen <italic>et al</italic> (<cite linkend="bb352493bib19" show="year">2005</cite>; rates of 10 and 30%), but we believe that the causes of these low rates differ between the studies.</p><p>The aerial trajectories of the fish in this study were much more consistent and accurately oriented relative to those observed in Alaska. Strongly variable mixed hydrodynamic signals from highly turbulent flows most likely did not exist in the OP, but they definitely did in the irregularly shaped pool below Brooks Falls. The low jump success rate observed in this study is more likely a result of the sharp right angle between the water channel and the vertical ramp in the AWG. This angle forced the water to separate immediately instead of flowing down the ramp for a short distance before separation, as was the case for both the rounded crest of Brooks Falls and the falls at the Russian River. Delaying the separation of water from the ramp with a rounded crest permitted the fish to push against water with their caudal fins when they landed directly on the crest. This resulted in successful jumps for many fish in Alaska. Without this water to push against, fish landing on the crest in this study were easily pushed back down into the pool.</p></sec-level2><sec-level2 id="bb352493s4-4" label="4.4"><heading><italic>Artificial waterfall generator</italic> design improvements</heading><p indent="no">Working with the AWG apparatus indicated several areas of desirable design improvement for future work.
<ordered-list id="bb352493ol2" pattern="2"><list-item id="bb352493ol2.1" marker="(i)"><p indent="no">One water pump provided ample power to drive the water delivery system for the fish tested. Fish congregated in the lower upstream corners of the OP, apparently avoiding the flow, during trials with one pump set at full throttle. They did not maintain those positions while both pumps were running.</p></list-item><list-item id="bb352493ol2.2" marker="(ii)"><p indent="no">A shallower maximum pool depth would allow for a reduction in the amount of steel reinforcement surrounding the system, which could reduce costs. It would also make possible a reduction in the required viewing panel area. Fish did not respond to the fall at any fall height (from 0 to 72 cm) when the pool depth was greater than 66 cm.</p></list-item><list-item id="bb352493ol2.3" marker="(iii)"><p indent="no">A rounded vertical fall crest is suggested. A crest without a radius promotes water flow separation much sooner than would a rounded crest. This prevents fish from successfully landing jumps at the crest.</p></list-item></ordered-list></p></sec-level2><sec-level2 id="bb352493s4-5" label="4.5"><heading>Conclusions and recommendations</heading><p indent="no">Sockeye salmon varieties, indeed probably most salmonid fishes, will jump up waterfalls in predictable ways when they are presented with suitable flow conditions. Flow rates must be strong enough to elicit rheotactic responses by the fish, but low enough to keep fish from retreating to regions protected from high flows (such as the corners of pools). These regions of lower flow are limited by using pool depths no deeper than the bottom of the boil formed by the falls. Preferred fall heights are dependent on the pool depth; mean depth/height ratios near 1.0 are optimal. However, flow rates must be taken into account. Gradients of falls have a strong influence on whether fish swim or jump up falls. A critical angle near 40° exists where the majority of attempts switch from swimming to jumping.</p><p>Jumping kinematics in salmonids differ from previously suggested mechanisms. Rather than passively rising to the surface and then accelerating using a C-start, fish initiate acceleration near the bottom of the pool, beneath the boil, with the more forward directed S-start followed by burst swimming to the surface. This jumping behavior is distinct from their behavior when swimming up falls at lower angles. Swimming approaches are initiated from the base of falls near the surface as opposed to beneath the boil. Consequently, pool and weir fish passageway designs must account for these differences by incorporating longer and deeper pools for jumping than are required for swimming.</p><p>The kokanee salmon of Taylor Creek demonstrate that populations of salmon isolated from the need to jump for many generations may still efficiently utilize pool and weir fish passageways. Quantitative differences between Alaskan sea run sockeye salmon and the kokanee used in this study indicate that fine tuning of designs for fish passageways probably will be needed for different populations. Adjustments are certain to be necessary for different species (Mallen-Cooper <cite linkend="bb352493bib21" show="year">1994</cite>, Peake <italic>et al</italic> <cite linkend="bb352493bib28" show="year">1997</cite>, Swanson <italic>et al</italic> <cite linkend="bb352493bib33" show="year">1998</cite>). The authors hope that application of the understanding of jumping behaviors described in this study may contribute to passageway designs that will better protect populations of threatened and endangered migratory stream fishes.</p><p>The preponderance of evidence appears to support the fact that complete removal of dams is the best way to restore populations of migratory fishes in river systems (Bowman <cite linkend="bb352493bib02" show="year">1996</cite>, Workman <cite linkend="bb352493bib41" show="year">2007</cite>). However, there are multiple very large dams on many rivers that are not likely to be removed in the foreseeable future for economic and political reasons. During the last 60+ years, fisheries biologists in many government agencies have introduced kokanee salmon into a wide range of freshwater habitats in North America (US Geological Survey Nonindigenous Aquatic Species Database, species no 915: <webref url="http://nas.er.usgs.gov/queries/FactSheet.aspx=speciesID=915">http://nas.er.usgs.gov/queries/FactSheet.aspx?speciesID=915</webref>; Seeley and McCammon <cite linkend="bb352493bib31" show="year">1966</cite>). Most of those introductions were made for reasons only marginally related to this paper. Many introductions put salmon into environments where they were completely exotic—actions that we consider problematic. However, the fact remains that a well-established exotic population is flourishing in the reservoir above the Flaming Gorge Dam in northeastern Utah (Colorado River drainage). Kokanee salmon, therefore, might be used as part of efforts at sustainable restoration ecology (Wilson <cite linkend="bb352493bib40" show="year">2002</cite>, Rosenzweig <cite linkend="bb352493bib29" show="year">2003</cite>) above dams where native sea run salmon populations have been extirpated from or are disappearing from suitable habitat upstream of the reservoirs impounded by dams that are unlikely to be removed.</p></sec-level2></sec-level1><acknowledgment><heading>Acknowledgments</heading><p indent="no">This study was funded by Sigma Xi Grants-in-Aid of Research (to DVL), the Department of Ecology and Evolutionary Biology at the University of California, Los Angeles, a grant from the University of California Centers for Water and Wildland Resources (project number W-928) and a grant from the Energy Science and Technology project of the UC Energy Institute (to MSG). We thank P B Moyle for insight into fisheries conservation issues. The engineering and developmental insights and overwhelming efforts in labor provided by T Lauritzen were crucial in the success of the AWG apparatus. We also thank J J Cech and W D Borgeson for their guidance during apparatus development. L H Caplan and A Caplan were extremely generous to allow crucial preliminary testing to be conducted in their private swimming pool. The computer aided design used in this manuscript was completed by T Lauritzen. We thank C Myrick for the initial suggestion of using kokanee salmon as study organisms. J Riener was extremely generous with his time by providing insights and site access to Taylor Creek. T Lauritzen, L H Caplan, W Quirk and P C O'Connor were instrumental in setting up experiments and data collection. 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Technol.</jnl-title><volume>24</volume><pages>31–42</pages></journal-ref></reference-list></references></back></article></istex:document></istex:metadataXml><mods version="3.6"><titleInfo lang="eng"><title>Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design</title></titleInfo><titleInfo type="abbreviated"><title>Salmon jumping</title></titleInfo><titleInfo type="alternative" lang="eng"><title>Salmon jumping: behavior, kinematics and optimal conditions, with possible implications for fish passageway design</title></titleInfo><name type="personal"><namePart type="given">D V</namePart><namePart type="family">Lauritzen</namePart><affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</affiliation><affiliation>E-mail: dlauritz@ccsf.edu</affiliation><role><roleTerm type="text">author</roleTerm></role></name><name type="personal"><namePart type="given">F S</namePart><namePart type="family">Hertel</namePart><affiliation>Department of Biology, California State UniversityNorthridge, Northridge, CA 91330, USA</affiliation><affiliation>E-mail: fritz.hertel@csun.edu</affiliation><role><roleTerm type="text">author</roleTerm></role></name><name type="personal"><namePart type="given">L K</namePart><namePart type="family">Jordan</namePart><affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</affiliation><affiliation>E-mail: msgordon@ucla.edu</affiliation><role><roleTerm type="text">author</roleTerm></role></name><name type="personal"><namePart type="given">M S</namePart><namePart type="family">Gordon</namePart><affiliation>Department of Ecology and Evolutionary Biology, University of California, Los Angeles, CA 90095-1606, USA</affiliation><affiliation>E-mail: ljordan@ucla.edu</affiliation><role><roleTerm type="text">author</roleTerm></role></name><typeOfResource>text</typeOfResource><genre type="research-article" displayLabel="paper">paper</genre><originInfo><publisher>IOP Publishing</publisher><dateIssued encoding="w3cdtf">2010</dateIssued><copyrightDate encoding="w3cdtf">2010</copyrightDate></originInfo><language><languageTerm type="code" authority="iso639-2b">eng</languageTerm><languageTerm type="code" authority="rfc3066">en</languageTerm></language><physicalDescription><internetMediaType>text/html</internetMediaType><note type="production">Printed in the UK</note></physicalDescription><abstract>Behavioral and kinematic properties and capacities of wild migratory salmonid fishes swimming upstream and jumping up waterfalls generally have played only minor roles in the design and construction of passageways intended to help these fishes get past dams and other human-made obstacles blocking their movements. This paper reports the results of an experimental study of relevant behavioral and kinematic properties of adult kokanee salmon (Oncorhynchus nerka) jumping up waterfalls as they migrate upstream. We used a portable, adjustable apparatus to study in the field fish responding to artificial waterfalls under a range of flow conditions. We observed fish under conditions of varying water flow rates, pool depths, fall heights and fall angles. We analyzed digital video recordings of their behaviors. Kokanee salmon spontaneously jump up waterfalls within a relatively narrow range of conditions, including low flow speeds, near vertical angles and pool depth to fall height ratios near 1.0. Preferred values for each parameter are, to some extent, dependent on other parameters. In contrast to previous misconceptions, jumping behavior is initiated by running S-start accelerations from beneath the boils formed in the plunge pools below waterfalls, as opposed to C-start standing jumps from the surface. S-starts are immediately followed by burst swimming to the point of takeoff at the surface. These results can contribute to an improved basis for developing designs of fish passageways that may ultimately make them more effective and efficient.</abstract><subject><genre>keywords</genre><topic>salmon</topic><topic>kokanee</topic><topic>jumping</topic><topic>waterfalls</topic><topic>behavior</topic><topic>kinematics</topic><topic>fish passageways</topic><topic>restoration ecology</topic></subject><relatedItem type="host"><titleInfo><title>Bioinspiration & Biomimetics</title></titleInfo><titleInfo type="abbreviated"><title>Bioinsp. Biomim.</title></titleInfo><genre type="journal">journal</genre><identifier type="ISSN">1748-3182</identifier><identifier type="PublisherID">bb</identifier><identifier type="CODEN">BBIICI</identifier><identifier type="URL">stacks.iop.org/BB</identifier><part><date>2010</date><detail type="volume"><caption>vol.</caption><number>5</number></detail><detail type="issue"><caption>no.</caption><number>3</number></detail><extent unit="pages"><start>1</start><end>11</end><total>11</total></extent></part></relatedItem><identifier type="istex">9ACE70A18185FE1AA829B2846A52A448DEC8EDA0</identifier><identifier type="DOI">10.1088/1748-3182/5/3/035006</identifier><identifier type="PII">S1748-3182(10)52493-1</identifier><identifier type="articleID">352493</identifier><identifier type="articleNumber">035006</identifier><accessCondition type="use and reproduction" contentType="copyright">2010 IOP Publishing Ltd</accessCondition><recordInfo><recordContentSource>IOP</recordContentSource><recordOrigin>2010 IOP Publishing Ltd</recordOrigin></recordInfo></mods></metadata><serie></serie></istex></record>Pour manipuler ce document sous Unix (Dilib)
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