http://lod.bco-dmo.org/id/dataset/773556
eng; USA
utf8
dataset
Highest level of data collection, from a common set of sensors or instrumentation, usually within the same research project
Biological and Chemical Oceanography Data Management Office (BCO-DMO)
Unavailable
508-289-2009
WHOI MS#36
Woods Hole
MA
02543
USA
info@bco-dmo.org
http://www.bco-dmo.org
Monday - Friday 8:00am - 5:00pm
For questions regarding this resource, please contact BCO-DMO via the email address provided.
pointOfContact
2019-07-22
ISO 19115-2 Geographic Information - Metadata - Part 2: Extensions for Imagery and Gridded Data
ISO 19115-2:2009(E)
Lab study on the effect of temperature and pCO2 on mussel byssal attachment (thread mechanics) with mussels collected in May 2012 from Argyle Creek, San Juan Island, WA (48.52˚ N, 123.01˚ W)
2019-07-24
publication
2019-07-24
revision
Marine Biological Laboratory/Woods Hole Oceanographic Institution Library (MBLWHOI DLA)
2019-08-06
publication
https://doi.org/10.1575/1912/bco-dmo.773556.1
Emily Carrington
University of Washington
principalInvestigator
Biological and Chemical Oceanography Data Management Office (BCO-DMO)
Unavailable
508-289-2009
WHOI MS#36
Woods Hole
MA
02543
USA
info@bco-dmo.org
http://www.bco-dmo.org
Monday - Friday 8:00am - 5:00pm
For questions regarding this resource, please contact BCO-DMO via the email address provided.
publisher
Cite this dataset as: Carrington, E. (2019) Lab study on the effect of temperature and pCO2 on mussel byssal attachment (thread mechanics) with mussels collected in May 2012 from Argyle Creek, San Juan Island, WA (48.52˚ N, 123.01˚ W). Biological and Chemical Oceanography Data Management Office (BCO-DMO). (Version 1) Version Date 2019-07-24 [if applicable, indicate subset used]. doi:10.1575/1912/bco-dmo.773556.1 [access date]
Dataset Description: <p>These data were used in a structural analysis study to evaluate how pCO2 and an additional stressor, elevated temperature, influences byssal thread quality and production. Mussels (<em>M. </em><em>trossulus</em>) were collected in May 2012 from Argyle Creek, San Juan Island, WA (48.52˚ N, 123.01˚ W) and held in a mesh box submerged under the dock at Friday Harbor Laboratories (FHL), San Juan Island, WA for up to 14 d. Mussels were placed in controlled temperature and pCO2 treatments in the Ocean Acidification Experimental Laboratory (OAEL), then newly produced threads were counted and pulled to failure to determine byssus strength.</p> Methods and Sampling: <p>Mussels (<em>M. trossulus</em>) were collected in May 2012 from Argyle Creek, San Juan Island, WA (48.52˚ N, 123.01˚ W) and held in a mesh box submerged under the dock at Friday Harbor Laboratories (FHL), San Juan Island, WA for up to 14 d. Mussels were placed in experimental mesocosms in the Ocean Acidification Experimental Laboratory (OAEL) at FHL as described in O’Donnell et al. (2013) and Timmins-Schiffman et al. (2012). Briefly, manipulations of pH were made by bubbling CO2 into a 150 L temperature-controlled seawater reservoir, that supplied water to eight 3.5 L chambers at a turnover rate of 50 ml min-1. Air was bubbled into the reservoir to maintain 100% oxygen saturation and submersible pumps (model number P396, Annex Depot, Sacramento, CA) provided mixing in the chambers at 3.8 L min-1. The bottom of each chamber was lined with autoclaved pebbles, collected from an FHL beach, to provide a substrate for byssal thread attachment. pH and temperature were monitored continuously in each water reservoir with a Durafet pH and temperature probe and the full carbonate chemistry of the system evaluated with DIC and Total alkalinity measurements once during each trial. Mussels were acclimated to their treatment temperatures in ambient pH (~7.8) over 9 d, ramping temperature up no more than 2˚C per day, and fed a maintenance level of Shellfish Diet 1800 (6 g l-1 day-1, Reed Mariculture, Campell, CA, USA).</p>
<p>The twelve independent temperature x pCO2 treatments spanned the range of local marine conditions (Newcomb, 2015; George <em>et al</em>., 2019; temperature at 10˚C, 18˚C, or 25˚C and pCO2 at 400, 750, 1200, or 2500 µatm). Each mussel was trimmed of external byssus before placement in an experimental treatment for 3 d, sufficient time to produce new mature byssal threads (Bell &amp; Gosline 1996) while minimizing the effect of treatment on mussel condition. Mussels were starved during the 3 d trials to minimize changes in chamber water chemistry due to food addition and to reduce fouling. Three trials were conducted in succession to replicate treatments over time, increasing sample size (n=8 x 3) for each temperature*pCO2 treatment.</p>
<p>At the end of each trial, mussels and the rocks to which they had attached with byssal threads were removed from the chambers. The entire byssus was dissected from each mussel and stored air-dried for up to 20 days. Byssus was rehydrated in seawater prior to testing, a method that does not alter the mechanical properties of the byssal threads (Brazee, 2004). The number of byssal threads each mussel produced was counted, and one thread was haphazardly chosen for mechanical testing following the procedure of Bell &amp; Gosline (1996). Briefly, an individual thread was clamped with submersible pneumatic grips on either end by holding the proximal byssal stem between cardstock with cyanoacrylate glue and affixing the distal plaque with attached rock to an aluminum T-bar with epoxy. An Instron 5565 tensometer (Norwood MA, USA), extended the thread at a rate of 10 mm min-1 in a temperature-controlled water bath (3130-100 BioPuls Bath, Instron, Norwood, MA, USA) until failure. The tensometer measured force (±10-3 N) and extension (± 10-3 mm) at 10 Hz. Tests were performed in seawater with a pH of 7.8 and the relevant treatment temperature.</p>
<p>Pull to failure mechanical tests provided estimates of thread breaking force, yield force, extensibility, initial stiffness and failure location (Bell &amp; Gosline 1996). Yield, due to quasi-plastic deformation in the distal region, was identified as the point where the initial slope of the force-extension curve decreased by 40%. Extensibility was calculated by dividing thread extension at failure by initial length and initial stiffness was determined from the initial slope of the force extension curve. The location of failure (proximal, plaque, and/or distal region) was noted and threads were retested to quantify the breaking force of each remaining region. Tests that broke at the grips were considered underestimates and were discarded.</p>
<p>The cross-sectional area of the proximal region was measured to evaluate morphological differences among treatments. The elliptical area was estimated from measures of the major and minor axes (+ 1 um using a dissecting microscope (Brazee &amp; Carrington 2006). Proximal breaking stress (N mm-2), a material property, was calculated as proximal breaking force divided by proximal area. Thread surface structure was examined using a scanning electron microscope (FEI Sirion XL30 SEM, Hillsboro, OR).</p>
<p>Whole mussel attachment strength was estimated using two mathematical models developed by Bell &amp; Gosline (1996). Each model assumes a mussel is anchored with a constant thread number (n=50) arranged in a circle. The normal model estimates dislodgment force perpendicular to the substrate (e.g, lift); all threads are engaged and extend until they reach their maximum force. The parallel model estimates dislodgement force for an animal pulled parallel to the substrate (e.g., drag); threads on the upstream side are the first in tension, yield and extend until they reach maximum force and break, while more threads are recruited into tension until they have all broken. Additionally, we modified each model to incorporate the variation in thread production across treatments. Because thread production was measured for only three days, treatment means were scaled to a maximum value of 50 threads.</p>
<p>Detailed methods and results are provided in Newcomb, 2015 and Newcomb <em>et al.,</em> 2019</p>
<p>Location: Friday Harbor Laboratories, Friday Harbor WA</p>
Funding provided by NSF Division of Ocean Sciences (NSF OCE) Award Number: OCE-1041213 Award URL: http://www.nsf.gov/awardsearch/showAward?AWD_ID=1041213
completed
Emily Carrington
University of Washington
206-221-4676
UW Friday Harbor Laboratories 620 University Road
Friday Harbor
WA
98250
USA
ecarring@uw.edu
pointOfContact
asNeeded
Dataset Version: 1
Unknown
Mussel_ID
Temp
pH
pCO2_Target
Treatment_ID
Trial
GI
CI
Distal_Diam
Prox_Major
Prox_Minor
Yield
Location
Breaking_Ext
Breaking_Force
Plaque_Force
Proximal_Force
Distal_Force
Breaking_Stress
Youngs_Mod
Thread_Stiffness
Thread_Ext
Instron 5565 load frame
theme
None, User defined
sample identification
temperature
pH
Partial pressure of CO2
treatment
experiment id
No BCO-DMO term
diameter
length
featureType
BCO-DMO Standard Parameters
Materials Testing System
instrument
BCO-DMO Standard Instruments
otherRestrictions
otherRestrictions
Access Constraints: none. Use Constraints: Please follow guidelines at: http://www.bco-dmo.org/terms-use Distribution liability: Under no circumstances shall BCO-DMO be liable for any direct, incidental, special, consequential, indirect, or punitive damages that result from the use of, or the inability to use, the materials in this data submission. If you are dissatisfied with any materials in this data submission your sole and exclusive remedy is to discontinue use.
Science, Engineering and Education for Sustainability NSF-Wide Investment (SEES): Ocean Acidification (formerly CRI-OA)
https://www.nsf.gov/funding/pgm_summ.jsp?pims_id=503477
Science, Engineering and Education for Sustainability NSF-Wide Investment (SEES): Ocean Acidification (formerly CRI-OA)
NSF Climate Research Investment (CRI) activities that were initiated in 2010 are now included under Science, Engineering and Education for Sustainability NSF-Wide Investment (SEES). SEES is a portfolio of activities that highlights NSF's unique role in helping society address the challenge(s) of achieving sustainability. Detailed information about the SEES program is available from NSF (https://www.nsf.gov/funding/pgm_summ.jsp?pims_id=504707).
In recognition of the need for basic research concerning the nature, extent and impact of ocean acidification on oceanic environments in the past, present and future, the goal of the SEES: OA program is to understand (a) the chemistry and physical chemistry of ocean acidification; (b) how ocean acidification interacts with processes at the organismal level; and (c) how the earth system history informs our understanding of the effects of ocean acidification on the present day and future ocean.
Solicitations issued under this program:NSF 10-530, FY 2010-FY2011NSF 12-500, FY 2012NSF 12-600, FY 2013NSF 13-586, FY 2014
NSF 13-586 was the final solicitation that will be released for this program.
PI Meetings:1st U.S. Ocean Acidification PI Meeting(March 22-24, 2011, Woods Hole, MA)2nd U.S. Ocean Acidification PI Meeting(Sept. 18-20, 2013, Washington, DC)
3rd U.S. Ocean Acidification PI Meeting (June 9-11, 2015, Woods Hole, MA – Tentative)
NSF media releases for the Ocean Acidification Program:
Press Release 10-186 NSF Awards Grants to Study Effects of Ocean Acidification
Discovery Blue Mussels "Hang On" Along Rocky Shores: For How Long?
Discovery nsf.gov - National Science Foundation (NSF) Discoveries - Trouble in Paradise: Ocean Acidification This Way Comes - US National Science Foundation (NSF)
Press Release 12-179 nsf.gov - National Science Foundation (NSF) News - Ocean Acidification: Finding New Answers Through National Science Foundation Research Grants - US National Science Foundation (NSF)
Press Release 13-102 World Oceans Month Brings Mixed News for Oysters
Press Release 13-108 nsf.gov - National Science Foundation (NSF) News - Natural Underwater Springs Show How Coral Reefs Respond to Ocean Acidification - US National Science Foundation (NSF)
Press Release 13-148 Ocean acidification: Making new discoveries through National Science Foundation research grants
Press Release 13-148 - Video nsf.gov - News - Video - NSF Ocean Sciences Division Director David Conover answers questions about ocean acidification. - US National Science Foundation (NSF)
Press Release 14-010 nsf.gov - National Science Foundation (NSF) News - Palau's coral reefs surprisingly resistant to ocean acidification - US National Science Foundation (NSF)
Press Release 14-116 nsf.gov - National Science Foundation (NSF) News - Ocean Acidification: NSF awards $11.4 million in new grants to study effects on marine ecosystems - US National Science Foundation (NSF)
SEES-OA
largerWorkCitation
program
Effects of Ocean Acidification on Coastal Organisms: An Ecomaterials Perspective
http://depts.washington.edu/fhl/oael.html
Effects of Ocean Acidification on Coastal Organisms: An Ecomaterials Perspective
<p><strong>Effects of Ocean Acidification on Coastal Organisms: An Ecomaterials Perspective</strong><br />
This award will support researchers based at the University of Washington's Friday Harbor Laboratories. The overall focus of the project is to determine how ocean acidification affects the integrity of biomaterials and how these effects in turn alter interactions among members of marine communities. The research plan emphasizes an ecomaterial approach; a team of biomaterials and ecomechanics experts will apply their unique perspective to detail how different combinations of environmental conditions affect the structural integrity and ecological performance of organisms. The study targets a diversity of ecologically important taxa, including bivalves, snails, crustaceans, and seaweeds, thereby providing insight into the range of possible biological responses to future changes in climate conditions. The proposal will enhance our understanding of the ecological consequences of climate change, a significant societal problem.</p>
<p>Each of the study systems has broader impacts in fields beyond ecomechanics. Engineers are particularly interested in biomaterials and in each system there are materials with commercial potential. The project will integrate research and education by supporting doctoral student dissertation research, providing undergraduate research opportunities via three training programs at FHL, and summer internships for talented high school students, recruited from the FHL Science Outreach Program. The participation of underrepresented groups will be broadened by actively recruiting URM and female students. Results will be disseminated in a variety of forums, including peer-reviewed scientific publications, undergraduate and graduate course material, service learning activities in K-8 classrooms, demonstrations at FHL's annual Open House, and columns for a popular science magazine.</p>
OA - Ecomaterials Perspective
largerWorkCitation
project
eng; USA
oceans
-123.01
-123.01
48.52
48.52
2012-05-01
2012-05-31
Friday Harbor, WA
0
BCO-DMO catalogue of parameters from Lab study on the effect of temperature and pCO2 on mussel byssal attachment (thread mechanics) with mussels collected in May 2012 from Argyle Creek, San Juan Island, WA (48.52˚ N, 123.01˚ W)
Biological and Chemical Oceanography Data Management Office (BCO-DMO)
Unavailable
508-289-2009
WHOI MS#36
Woods Hole
MA
02543
USA
info@bco-dmo.org
http://www.bco-dmo.org
Monday - Friday 8:00am - 5:00pm
For questions regarding this resource, please contact BCO-DMO via the email address provided.
pointOfContact
http://lod.bco-dmo.org/id/dataset-parameter/773600.rdf
Name: Mussel_ID
Units: unitless
Description: Mussel sample identifier
http://lod.bco-dmo.org/id/dataset-parameter/773601.rdf
Name: Temp
Units: degrees Celcius (°C)
Description: Temperature
http://lod.bco-dmo.org/id/dataset-parameter/773602.rdf
Name: pH
Units: unitless
Description: pH (total scale)
http://lod.bco-dmo.org/id/dataset-parameter/773603.rdf
Name: pCO2_Target
Units: microatmosphere (uatm)
Description: Target pCO2 level
http://lod.bco-dmo.org/id/dataset-parameter/773604.rdf
Name: Treatment_ID
Units: unitless
Description: Treatment identifier
http://lod.bco-dmo.org/id/dataset-parameter/773605.rdf
Name: Trial
Units: unitless
Description: Trial identifier
http://lod.bco-dmo.org/id/dataset-parameter/773606.rdf
Name: GI
Units: unitless
Description: Gonad Index
http://lod.bco-dmo.org/id/dataset-parameter/773607.rdf
Name: CI
Units: gram per cubic millimeter (g/mm^3)
Description: Condition Index
http://lod.bco-dmo.org/id/dataset-parameter/773608.rdf
Name: Distal_Diam
Units: millimeter (mm)
Description: Thread distal region diameter
http://lod.bco-dmo.org/id/dataset-parameter/773609.rdf
Name: Prox_Major
Units: millimeter (mm)
Description: Thread proximal region major axis length
http://lod.bco-dmo.org/id/dataset-parameter/773610.rdf
Name: Prox_Minor
Units: millimeter (mm)
Description: Thread proximal region minor axis length
http://lod.bco-dmo.org/id/dataset-parameter/773611.rdf
Name: Yield
Units: Newton (N)
Description: Thread yield force
http://lod.bco-dmo.org/id/dataset-parameter/773612.rdf
Name: Location
Units: unitless
Description: Thread failure location
http://lod.bco-dmo.org/id/dataset-parameter/773613.rdf
Name: Breaking_Ext
Units: millimeter (mm)
Description: Thread breaking extension
http://lod.bco-dmo.org/id/dataset-parameter/773614.rdf
Name: Breaking_Force
Units: Newton (N)
Description: Thread breaking force
http://lod.bco-dmo.org/id/dataset-parameter/773615.rdf
Name: Plaque_Force
Units: Newton (N)
Description: Thread plaque breaking force
http://lod.bco-dmo.org/id/dataset-parameter/773616.rdf
Name: Proximal_Force
Units: Newton (N)
Description: Thread proximal region breaking force
http://lod.bco-dmo.org/id/dataset-parameter/773617.rdf
Name: Distal_Force
Units: Newton (N)
Description: Thread distal region breaking force
http://lod.bco-dmo.org/id/dataset-parameter/773618.rdf
Name: Breaking_Stress
Units: Newton per square millimeter (N/mm^2)
Description: Thread breaking strength
http://lod.bco-dmo.org/id/dataset-parameter/773619.rdf
Name: Youngs_Mod
Units: Newton per square millimeter (N/mm^2)
Description: Thread Youngs modulus
http://lod.bco-dmo.org/id/dataset-parameter/773620.rdf
Name: Thread_Stiffness
Units: Newton per square millimeter (N/mm^2)
Description: Thread stiffness
http://lod.bco-dmo.org/id/dataset-parameter/773621.rdf
Name: Thread_Ext
Units: unitless
Description: Thread extensibility
GB/NERC/BODC > British Oceanographic Data Centre, Natural Environment Research Council, United Kingdom
Biological and Chemical Oceanography Data Management Office (BCO-DMO)
Unavailable
508-289-2009
WHOI MS#36
Woods Hole
MA
02543
USA
info@bco-dmo.org
http://www.bco-dmo.org
Monday - Friday 8:00am - 5:00pm
For questions regarding this resource, please contact BCO-DMO via the email address provided.
pointOfContact
24078
https://darchive.mblwhoilibrary.org/bitstream/1912/24431/1/dataset-773556_thread-mechanics__v1.tsv
download
https://doi.org/10.1575/1912/bco-dmo.773556.1
download
onLine
dataset
<p>Mussels (<em>M. trossulus</em>) were collected in May 2012 from Argyle Creek, San Juan Island, WA (48.52˚ N, 123.01˚ W) and held in a mesh box submerged under the dock at Friday Harbor Laboratories (FHL), San Juan Island, WA for up to 14 d. Mussels were placed in experimental mesocosms in the Ocean Acidification Experimental Laboratory (OAEL) at FHL as described in O’Donnell et al. (2013) and Timmins-Schiffman et al. (2012). Briefly, manipulations of pH were made by bubbling CO2 into a 150 L temperature-controlled seawater reservoir, that supplied water to eight 3.5 L chambers at a turnover rate of 50 ml min-1. Air was bubbled into the reservoir to maintain 100% oxygen saturation and submersible pumps (model number P396, Annex Depot, Sacramento, CA) provided mixing in the chambers at 3.8 L min-1. The bottom of each chamber was lined with autoclaved pebbles, collected from an FHL beach, to provide a substrate for byssal thread attachment. pH and temperature were monitored continuously in each water reservoir with a Durafet pH and temperature probe and the full carbonate chemistry of the system evaluated with DIC and Total alkalinity measurements once during each trial. Mussels were acclimated to their treatment temperatures in ambient pH (~7.8) over 9 d, ramping temperature up no more than 2˚C per day, and fed a maintenance level of Shellfish Diet 1800 (6 g l-1 day-1, Reed Mariculture, Campell, CA, USA).</p>
<p>The twelve independent temperature x pCO2 treatments spanned the range of local marine conditions (Newcomb, 2015; George <em>et al</em>., 2019; temperature at 10˚C, 18˚C, or 25˚C and pCO2 at 400, 750, 1200, or 2500 µatm). Each mussel was trimmed of external byssus before placement in an experimental treatment for 3 d, sufficient time to produce new mature byssal threads (Bell &amp; Gosline 1996) while minimizing the effect of treatment on mussel condition. Mussels were starved during the 3 d trials to minimize changes in chamber water chemistry due to food addition and to reduce fouling. Three trials were conducted in succession to replicate treatments over time, increasing sample size (n=8 x 3) for each temperature*pCO2 treatment.</p>
<p>At the end of each trial, mussels and the rocks to which they had attached with byssal threads were removed from the chambers. The entire byssus was dissected from each mussel and stored air-dried for up to 20 days. Byssus was rehydrated in seawater prior to testing, a method that does not alter the mechanical properties of the byssal threads (Brazee, 2004). The number of byssal threads each mussel produced was counted, and one thread was haphazardly chosen for mechanical testing following the procedure of Bell &amp; Gosline (1996). Briefly, an individual thread was clamped with submersible pneumatic grips on either end by holding the proximal byssal stem between cardstock with cyanoacrylate glue and affixing the distal plaque with attached rock to an aluminum T-bar with epoxy. An Instron 5565 tensometer (Norwood MA, USA), extended the thread at a rate of 10 mm min-1 in a temperature-controlled water bath (3130-100 BioPuls Bath, Instron, Norwood, MA, USA) until failure. The tensometer measured force (±10-3 N) and extension (± 10-3 mm) at 10 Hz. Tests were performed in seawater with a pH of 7.8 and the relevant treatment temperature.</p>
<p>Pull to failure mechanical tests provided estimates of thread breaking force, yield force, extensibility, initial stiffness and failure location (Bell &amp; Gosline 1996). Yield, due to quasi-plastic deformation in the distal region, was identified as the point where the initial slope of the force-extension curve decreased by 40%. Extensibility was calculated by dividing thread extension at failure by initial length and initial stiffness was determined from the initial slope of the force extension curve. The location of failure (proximal, plaque, and/or distal region) was noted and threads were retested to quantify the breaking force of each remaining region. Tests that broke at the grips were considered underestimates and were discarded.</p>
<p>The cross-sectional area of the proximal region was measured to evaluate morphological differences among treatments. The elliptical area was estimated from measures of the major and minor axes (+ 1 um using a dissecting microscope (Brazee &amp; Carrington 2006). Proximal breaking stress (N mm-2), a material property, was calculated as proximal breaking force divided by proximal area. Thread surface structure was examined using a scanning electron microscope (FEI Sirion XL30 SEM, Hillsboro, OR).</p>
<p>Whole mussel attachment strength was estimated using two mathematical models developed by Bell &amp; Gosline (1996). Each model assumes a mussel is anchored with a constant thread number (n=50) arranged in a circle. The normal model estimates dislodgment force perpendicular to the substrate (e.g, lift); all threads are engaged and extend until they reach their maximum force. The parallel model estimates dislodgement force for an animal pulled parallel to the substrate (e.g., drag); threads on the upstream side are the first in tension, yield and extend until they reach maximum force and break, while more threads are recruited into tension until they have all broken. Additionally, we modified each model to incorporate the variation in thread production across treatments. Because thread production was measured for only three days, treatment means were scaled to a maximum value of 50 threads.</p>
<p>Detailed methods and results are provided in Newcomb, 2015 and Newcomb <em>et al.,</em> 2019</p>
<p>Location: Friday Harbor Laboratories, Friday Harbor WA</p>
Specified by the Principal Investigator(s)
<p>BCO-DMO processing notes:</p>
<ul>
<li>Renaming column headers</li>
</ul>
Specified by the Principal Investigator(s)
asNeeded
7.x-1.1
Biological and Chemical Oceanography Data Management Office (BCO-DMO)
Unavailable
508-289-2009
WHOI MS#36
Woods Hole
MA
02543
USA
info@bco-dmo.org
http://www.bco-dmo.org
Monday - Friday 8:00am - 5:00pm
For questions regarding this resource, please contact BCO-DMO via the email address provided.
pointOfContact
Instron 5565 load frame
Instron 5565 load frame
PI Supplied Instrument Name: Instron 5565 load frame PI Supplied Instrument Description:Instron’s (Norwood, MA) electromechanical testing systems are used to test a wide range of materials in tension or compression. The series 5560 are dual column table top models, the 5565 model has a load capacity of 5 kN (1125 lbf). Instrument Name: Materials Testing System Instrument Short Name:Materials Testing System Instrument Description: Testing systems that are used to test a wide range of
materials in tension or compression.