vvEPA
            United States
            Environmental Protection
            Agency
            Environmental Research
            Laboratory
            Duluth MN 55804
EPA/600/3-87/001
January 1987
            Research and Development
Guidelines
for the Culture of
Fathead Minnows
Pimephales
Prom el as for Use in
Toxicity Tests

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                                         EPA/600/3-87/001
                                         January 1937
GUIDELINES FOR THE CULTURE OF FATHEAD MINNOWS
PIMEPHALES PROMELAS FOR USE IN TOXICITY TESTS
                      by

               Jeffrey  S.  Denny
 Environmental Research Laboratory  -  Duluth
            Duluth, Minnesota 55804
 ENVIRONMENTAL RESEARCH LABORATORY  -  DULUTH
     OFFICE OF RESEARCH AND DEVELOPMENT
    U.S.  ENVIRONMENTAL PROTECTION AGENCY
            DULUTH, MINNESOTA 55804
                          U.S. Environments! Faction Agency
                          D^^'"i.. P-, i ;ur. - •  f<~>i  ion
                          77 V'i'\i Jackson Buu c.ard, 12th Floor
                          Chicago, IL 60604-3590

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                      NOTICE

This document has been reviewed in accordance with
U.S. Environmental Protection Agency policy and
approved for publication.  Mention of trade names
or commercial products does not constitute endorse-
ment or recommendation for use.

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                          FOREWORD

     Fathead minnows,  Pimephales promelas Rafinesque,  have  been
cultured at the Environmental Research Laboratory - Duluth for
use in aquatic toxicity  tests since the establishment of the
laboratory in 1967.   The  techniques and  apparatus described in
this report were developed over the years  by many  researchers.
     This paper sets  forth  the conditions and procedures now
being used to  produce research quality fathead minnow embryos,
larvae, juveniles, and adults. These guidelines can be modified
to adapt to  different circumstances  and needs.

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                           ABSTRACT



     This  paper  describes  the mechanical  apparatus and



biological  techniques now  in use to culture  fathead minnows



                             the us  Environmental  Protection
Agency's  Environmental Research Laboratory in Duluth,  Minnesota.



Physical  system information includes  water supply,  construction



materials,  water temperature, photoperiod,  and the water delivery



system.  The  biological  section addresses the  selection  of



spawning fish,  incubation of embryos, larval and adult feeding,



disease,  and gene pool considerations. This document is meant to



be a guide for those interested in culturing fathead minnows for



use in fish toxicology research.

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                       ACKNOWLEDGEMENTS







     For continual help throughout  the  development  of this guide



I would like to thank Larry Herman, Barb Halligan,  Jodi Collins,



and Andy Peterson.  For expert advice on the  culture of fatheads



I would like to thank Barb Halligan, Armond Lemke,  Duane Benoit,



Dick Carlson,  and  Don Mount.  For manuscript review I am indebted



to all  of  the above mentioned people,  as well  as  to Richard



Siefert,  Quentin Pickering,  J. Howard McCormick,  Teresa Norberg-



King,  Scott Heinritz,  John Brazner,  and Jim Taraldsen.   I extend



special  thanks to Jodi  Collins  for   illustrations  and  Eric



Peterson for photographic  support.



     Much of the credit for this document is due to present and



former staff members of the Environmental Research Laboratory  -



Duluth,  who have,  over  the years, developed this  system of



fathead minnow culture.

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                            CONTENTS

Foreword                                                  ii
Abstract                                                 iii
Acknowledgements                                          iv

1.  Purpose                                                 1

2.  Design                                                  1

3.  Physical System

     3.1   Tanks                                           3
     3.2   Water supply                                    4
     3.3   Water delivery system                           6
     3.4   Construction materials                          9
     3.5   Temperature                                    10
     3.6   Aeration                                       10
     3.7   Photoperiod                                    11
     3.8   Spawning substrates                            11
     3.9   Fail safe devices                              12
     3.10  Tank cleaning                                  12
     3.11  Disturbance                                    14
     3.12  Brine shrimp hatchery                          14

4.  Biological System

     4.1   Obtaining brood stock                          16
     4.2   Selection of spawning fish                     17
     4.3   Spawning                                       21
     4.4   Embryo incubation                              23
     4.5   Larval feeding                                 27
     4.6   Adult feeding                                  29
     4.7   Disease                                        29
     4.8   Gene pool                                      31
     4.9   Record keeping                                 32

5.  Literature cited                                       33

6.  Additional reading                                     35

7.  List of suppliers                                      41
                               vn

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                           PURPOSE



     A fathead minnow  culture  facility provides a continuous



supply of embryos  or fish of known  age,   raised under known



conditions, for aquatic toxicity testing. The use of laboratory



reared animals is advantageous since age and genetic background



are known, diet is controlled,  fish are free  from disease, and



are available  year-round.



     This document describes in detail fathead minnow culture



techniques used  at the Environmental Protection Agency's



Environmental  Research Laboratory  in Duluth, Minnesota.   Present



practices are based on years of  expertise developed by staff



scientists.  Local  conditions and specific needs of  researchers



may require modification of these guidelines.








                           DESIGN







     The  life  stages of fish in greatest demand for testing are



less than 24 hr old embryos, 0 - 24 hr old larvae, and 30 day old



juveniles. This system is designed to best produce these three



life stages.



     The  fathead culture unit at ERL-D has  54 m2 of floor  space



in  two  adjacent rooms.   The main room  (36  m2) contains 24



spawning  tanks,  60  juvenile rearing tanks,  20  brood stock tanks,



6  hatching  trays  for  eggs,   and  20  more tanks  for



experimental/emergency use (See Photograph #1).  This provides



ample space for 96 pairs  of adult spawners, 400 - 500 maturing

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I
42
-S

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fish for use as future spawning stock,  and about 15,000 juveniles
being reared to 30 days of age.  This system produces from 1000
- 2000 eggs/day,  and provides  400  -  500  30 day old juveniles
each day for toxicity  testing.    The  adjacent room  (18 m2)
provides space to quarantine and  breed  incoming wild fish used to
intermittently diversify the gene pool.
     Production now supplies about 15 - 20 researchers engaged in
full time toxicity testing ,  and fills occasional requests from
other scientists.  A slight  over -  production is necessary to
allow for variations in egg production and for recycling spare
fish  into new brood stock.
                        PHYSICAL SYSTEM

3.1  Tanks

     The tanks  are 57 liter (15 gallon) glass aquaria,  31  cm x  61
cm x 32 cm deep, with standpipe drains adjusted to provide  20  cm
of water depth.  These contain approximately  40  liters of  water
per tank.  The tanks, supported on racks made of slotted  angle
iron and 1.9 cm (3/4  inch) plywood,  are  arranged in 2 tiers,  in
rows of 12 tanks.  Two rows placed back to back form a bank of  48
tanks.   Three such banks make up the entire culture system.
     When starting a  culture program,  new tanks must be acid (10%
nitric), and acetone rinsed  before using.   Used tanks must  be
disinfected  with hypochlorite.  Allow  culture water  to flow

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through  the tanks for a day or  two before adding fish.
     For spawning fish, divide  the tanks   into four sections
with stainless  steel  screen (eg:  5  mm mesh,  0.89  mm  wire), glued
in place with  silicon glue.   The  screen  material must be acid
leached  before  use to  remove machine oil.   A single  water inlet,
drain,  and air  stone  can service each tank.
     White plastic dishpans,  commonly available in department
stores  (eg:  53 cm x 40 cm x 12 cm deep)  make good hatching trays
for embryo incubation.   White  trays provide the  best  background
color for seeing the newly hatched larvae.  Place six hatching
trays in a temperature controlled water bath (eg: 130 cm x 125 cm
x  8  cm  deep,  constructed from 1.9 cm (3/4  ijich)  plywood and
sealed on the inside with epoxy paint).  If this bath is placed in
a rack below a bank  of fish tanks,  drain water from the fish
tanks will  provide an inexpensive heating  source for embryo
hatching.  Fish tanks at 25°C will  warm  hatching  pans to 23°C.
3.2  Water Supply


     Unfiltered  Lake  Superior water is the water supply for  the
ERL - D fathead culture unit.   It has a pH range of 7.4 - 8.2,
alkalinity (as CaCO3) of  42 mg/1,  and  total hardness of 45 mg/1.
For more  detailed chemical  characterization of Lake  Superior
water, see Glass (1977).

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     Water supply must be the most important consideration in
establishing a fathead culture facility.  Use a natural supply
such  as a spring,  well, or  controlled  surface  water with
consistent water quality, if possible.    Culture water should be
similar to water  used in  testing.   Examine  the source  for
contamination  by  pesticides,  heavy metals, sulfides,  disease
vectors,  or any other suspected contaminants.   Make checks of
water quality periodically.   Filtration may be necessary if well
water  is  used  (Mount,  1971).   Dechlorinated tap water  from a
municipal water  supply should be  used  only as a  last  resort
(Benoit, 1982).  If  the water supply is  contaminated with fish
pathogens, pass the water  through an ultraviolet or  similar
sterilizer immediately  before  it enters  the system  (Allison and
Hermanutz, 1977).
     Water quality  parameters  such  as hardness,  alkalinity,  and
anions should fall  within the  following  limits:  hardness - 40 -
300 mg/1  (as  CaCO3),  and  alkalinity slightly less than  the
hardness.  The anions should be those found  in a normal stream or
lake.  Avoid well or spring waters which  have high iron, silica,
sulfides,  or chlorides not found in  surface waters.
     The quantity  of water necessary depends on the size of the
intended  culture unit.   The ERL  - D system of  over 150 tanks
consumes 15 - 20  liters/minute  when in full operation.  Smaller
systems,  or  systems in areas of  limited water supply could
operate on a reduced flow.  Though less desirable, static renewal
or recirculating systems may be effective.

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3.3   Water Delivery System

     The  ERL - D water delivery  system is a constant temperature,
flow-through  system.   It is gravity fed,  with custom  welded
stainless steel  mixing  boxes  (46 cm  x  28 cm  x 40 cm deep)
positioned in an open ceiling approximately 3.5  m  overhead.  Lake
Superior  water warmed  to  20°C flows through a toilet tank valve1
to maintain water level  in the headbox.   Water heated to 30°C
flows through  a solenoid valve2 positioned over the headbox.  To
achieve the desired constant temperature, a temperature probe3
suspended in  the  headbox  controls  a  solid state temperature
controller (Syrett and Dawson,  1975). This controller opens and
closes the solenoid valve when resistance  of the temperature
probe changes due to  change in temperature. When the  headbox
temperature  falls below  25°,  the solenoid valve   activates,
adding 30° water until  the headbox temperature  reaches 25°,  when
the  probe/relay   system  closes the solenoid  valve.   Headbox
temperature  must  be maintained at 26-  27° to provide 25° at the
tanks.  This is due to heat loss to ambient temperature.  This
basic water delivery system can  supply most experimental water

     ^ Numbers refer to suppliers that  manufacture or distribute
the type of product described.  A list of suppliers is appended.
Mention of trade names or commercial products does not constitute
endorsement  or recommendation for  use  by  the U.S. Environmental
Protection  Agency.

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temperatures, depending only on the temperature of the water



supply to  the  float and solenoid  valves.  (See Figure  #1).



     Air stones  or some  other type of agitator4 must be used in



the headbox to assure complete  mixing and oxygenation, and to



prevent supersaturation of gases  caused by heating  water.  Water



flows from the headbox through a 1.27 cm (1/2 inch) threadable,



polyvinyl  chloride  (PVC) pipe  (schedule  80,  available  from



plumbing  supply houses).  The  outlet is on  the side of the



headbox, about 2 -3  cm up from the bottom. Detailed instructions



on the design of the mixing boxes, electronic relays, etc., are



available in Syrett and Dawson, 1972, 1975, and McCormick and



Syrett,  ms,  1970.



     Water flows into the tanks  through a 1.27 cm  (1/2 inch) PVC



pipe manifold.  Above each tank there  is a T with  a 1.27 cm  (1/2



inch) to .95 cm  (3/8 inch) reducer attached.  Into the reducer  a



3 ml  disposable syringe barrel5 is glued with silicone glue.



This allows  the  use of different  sizes of hypodermic needles5 to



control flow rates.   Seventeen gauge needles provide 100 - 150



ml/min with  1 - 2 m head pressure.

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3.4  Construction Materials

     Construction materials  which come  in contact with the water
must not contain leachable substances.  Rubber,  copper, brass,  or
plastics  containing  fillers,  additives,  stabilizers,
plasticizers, etc,  must not  be used (Mount,1971).   Glass,
stainless steel, teflon,and PVC are the preferred construction
materials.  All  piping should be  of  rigid  PVC.  Schedule 80,
threadab^e  pvc  must be used to  avoid the danger of  toxicity
from PVC glue.  Silicone6 glue  is safe to use as long as enough
curing time is allowed.   (Follow  manufacturers instructions for
complete curing.)  Check all batches  of neoprene stoppers for
toxicity prior to use.  Recent  static tests at ERL - D show that
certain types  of neoprene stoppers  are acutely toxic to  fathead
minnow larvae.   A simple 24 hr  static test exposing fathead
larvae to the  stoppers in a beaker of culture  water will indicate
whether the stoppers  are safe.
     Ground fault interrupters7   are necessary  on all electrical
components  because  of the  close proximity  of electricity and
water  in these  systems.  Electrical  equipment must  be three
pronged and well grounded.  Avoid hanging extension cords,  or
extension cords on the floor,  where tank  overflow could cause
them to get wet.

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3.5  Temperature







     Water temperature in  all  tanks in  the  ERL -  D  fathead



culture  unit is maintained at 25°C  +  1°C   for  maximum  egg



production and growth.  Brungs (1971a) found that temperatures



below 22°C or above 26°C reduced fathead  minnow reproduction.



The described temperature control system  will maintain 25  +  i°c.



Culture  systems will function down to 22°C.







3.6  Aeration







     Provide  continuous gentle aeration to the tanks  to maintain



dissolved oxygen  concentrations above 5.0 mg/1  at all  times,  but



avoid vigorous aeration, especially with newly hatched larvae.



If a level of 5.0 mg/1 cannot be maintained,  remove some fish



from the tank (Mount, 1971).  Brungs (1971b),  found that larval



growth was the most sensitive indicator  of sublethal  effects of



lowered dissolved oxygen, and significant effects were seen at. 5



mg/1.  Provide an oil  free air supply of  at least 150  cc/min per



tank.  An oil trap or filter may be necessary on some systems.



Check the location of air intakes  and  efficient  operation of



laboratory air compressors to avoid  introducing contaminants.



     Arrange air tubing,  brass air valves,  and  air stones,



(obtainable  in aquarium stores) to provide an airline to each



tank.   Incorporate  a pressure regulator into the  system and feed



each group of  6  or 12 tanks  from  a central, larger diameter





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manifold. A pressure regulator on the main manifold will allow
use of valves with slip-on tubing,  without blowing the tubing
off.

3.7  Photoperiod

     Use  lights that simulate the wavelength spectra of sunlight.
A combination of  Durotest Optima FS8 and wide spectrum Sylvania
Gro - Lux9  fluorescent tubes has proven satisfactory in the  ERL -
D system.   Light  intensities at the  water surface should average
400 - 500 lux.    The photoperiod should be constant at 16 hours
light/8 hours  dark (Mount, 1971).   Gradual  changes in light
intensity at dawn and dusk may  be included within the photoperiod
if desired (see Drummond and Dawson, 1970).  Paragon Model  #4001
timers10  control photoperiod  in  the ERL -  D  system.  Lighting
warms the culture water, and this effect must be considered as
part of the overall design.

3.8  Spawning Substrates

     Since fathead minnows deposit their  adhesive eggs on the
underside of submerged or floating objects, the  spawning
substrate provided must yield a  similar surface.   The  ERL - D
culture system uses fiber cement water pipe,  7.6 to 10.2  cm in
diameter, cut  into 7-10 cm long  sections.   When halved
lengthwise and  inverted, these  pipe  sections form a semicircular

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arch.  Benoit (1982),  used  stainless steel  (16 - 18 gauge), cut



and bent into the same size and shape with a thin layer of quartz



sand glued on the underside with silicone glue.    This type of



substrate can come unglued after prolonged use, however.  Gale



and Bunyak  (1982) ,  successfully used   PVC  pipe  in the  same



configuration.  (See Photograph #2 for general view of spawning



and egg incubation  apparatus).







3.9  Fail  Safe Devices







     An alarm  system or fail-safe device will protect  against



temperature control or  water flow failure.   Temperature



recorders11  monitored hourly by security personnel  are in use at



ERL - D. Alternatively, use high - low  alarm systems,  or solenoid



valves to shut off water to the  tanks if the temperature rises



above 28°  or falls below 20°C.







3.10  Tank Cleaning







     ERL - D personnel  scrape  tanks  and siphon  residue  on a



biweekly,  rotating schedule. Some tanks are siphoned between



cleanings if necessary.  Most  culturists  allow light growths of



algae,  rotifers,  etc, to  remain,  for these provide a  dietary



supplement  for  fish.  Excessive  growths  of blue-green algae or



fungus must  be removed. Test cleaning tools for possible toxicity



before use.





                             12

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3.11   Disturbance
           *


     Shield the fish from  continual or drastic disturbance. Avoid

construction noises, continual human presence, and  extraneous

lights that might alter the photoperiod.



3.12   Brine Shrimp (Artemia spp.)   Hatchery



     The brine shrimp hatchery at ERL  - D  consists  of a

rectangular fiberglass tank, 2.1 m  x 0.6 m x 0.5 m deep, that

serves as a water bath.  (See Photograph  #3).  Rails mounted on

the inside,  9 cm down from the top, support 10 to 12 movable

plywood racks  (53 cm x 15 cm).   Each of these racks has three

11.5  cm diameter holes that hold  brine shrimp hatching jars.,  A

manifold  of  airline  tubing  provides an  air supply  to each

hatching jar.

     Brine shrimp will hatch in any container  with a conical

bottom, as long as  air bubbled  at the bottom of the cone keeps

the eggs  in  constant motion.  Some  culturists  use separatory

funnels, and others use inverted 1  liter plastic soda bottles

with the bottoms cut out.

     The hatching jars in use  at ERL -  D  are made from round

bottom glass  light  fixtures12, about  11 cm in diameter  and 24 cm

long.  A ring of airline tubing  glued around the jar  5 cm from

the open end suspends  each jar  through  a  hole in the plywood
                             14

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15

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rack,  partially submerged in the water bath. The capacity of each



jar is  about 1.5 liters.  This system provides  for up  to  36



hatching jars. A temperature  probe and controller system similar



to the headboxes  feeding the fish tanks maintains the bath  at



25°C.
                       BIOLOGICAL SYSTEM







4.l  Obtaining  Brood Stock







     Fish that  are  free of disease  and  adapted to laboratory



conditions make the best initial brood stock.  For the least risk



of disease,  and greater ease of shipment, begin with embryos.



Use of embryos also avoids any bioaccumulation of toxicants that



may  occur with adults.   Less  desirable is  the  use of  fish



captured in the wild or purchased from  a  bait dealer.  Take care



to verify that the animals are PAniejDliaJ.es^ p_romej.as^  and not a



related species.  Examine all fish,  and especially   wild caught



or bait  dealer fish,  for  signs  of  disease.   Use prophylactic



treatment (described in  section 4.7)   to insure good health.



These fish should then be bred through one full generation to



determine vigor, fecundity, and freedom from disease before use



in toxicity testing.
                              16

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4.2  Selection of spawning fish




     Determination of the  sex  of  immature fathead minnows is

nearly impossible, which makes it difficult to select mating

pairs before the fish actually begin breeding.  Breeding males

develop  a conspicuous gray pad of spongy  tubercles on the dorsal

surface  anterior to  the dorsal  fin, and two  rows of strong

tubercles across the snout.   The sides of the body become almost

black except  for two wide vertical bars which are  light  colored.

Another  characteristic of the breeding male is the presence  of a

dark spot at  the  anterior insertion of the dorsal fin.    Females

remain quite  drab (Eddy and Underbill, 1974).  The female fathead

minnow exhibits an ovipositor at least a month before spawning

(Flickinger,  1969).  (See  Photographs #4 & 5,  and Figure  #2).

     Breeder fish are selected as follows:  juvenile fish 3-4

months old are stocked at a density of 30 -  40 fish per 15 gallon
                                                 T
tank,  and provided with  2 or  3  spawning substrates as previously

described.   The presence of spawning substrates   hastens the

maturation process.  In 1 -  2 weeks some males  in the tank will

show  signs  of maturing.  Females will  begin to become gravid

soon after the males exhibit spawning color. For  observation,

net fish  from the tank and place individually  into a 400 ml

beaker with  approximately 3  cm of water.  Sexual maturity can

then be  determined as  previously  described.  Backlighting  makes

the female ovipositor easier to see.

     Removal of the mature fish for service as spawners  will



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                       •8*


                       8
                        &1 (0
                        H & *



                        ffl 

                       fe ft (0
20

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stimulate  the  subordinate juveniles in the brood  stock tanks to



ripen,  and take the places of the dominant  fish that were removed



from the hierarchy.  This method will provide a continuous source



of mature fish.   Some of  each month's leftover larvae or



juveniles not used  in testing  must be set up in  brood stock



tanks to begin the maturation process.  The ERL -  D system uses



16 brood tanks,  along with 8 larger  holding tanks (208 liters)



for bringing fish to maturity.







4.3  Spawning







     The fathead minnow is an intermittent, multiple spawning



species with  an extended breeding  season,  possibly spawning



intermittently all summer (Hasler,  1946; Radcliff,  1931).  Under



controlled culture conditions  fathead minnows will spawn



throughout the year.



     Fathead minnows usually spawn beneath objects, even under



such transient objects as  old maple and oak leaves (Isaak, 1961).



Males clean the underside of the  object  selected for the nest



site using the head tubercles in a scraping action and pulling



pieces of  algae  and associated debris off  the nest surface with



the mouth (Andrews and Flickinger, 1973).  The male and female



swim back and  forth beneath  the prepared overhead site and  roll



on their sides to  emit the sex products.   Close lateral contact



and body vibration characterize the act of spawning.  McMillan



(1972)  describes the spawning act as follows: "Finally, when a





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sufficient  degree of vibratory stimulation has been reached, the



male lifts and presses the female's  ventral surface against the



object's underside. In doing  so, he turns so that he is beneath



the  female and  can use  the posterior part  of his body to



manipulate  her upward.  The tubercules on his large pectoral fins



also help him  to  grip the  female tightly.   As the fishes' bodies



are taut and strained in this position, the female emits one or



perhaps several eggs,  and the male probably  releases sperm at



this instant.  Then they abruptly separate, although a new bout



of vibrating may begin  only seconds later."



     The buoyant,  adhesive  embryos stick to each  other  and to the



undersurface  of  the nesting  object.   After    deposition is



complete,  the  male  remains at the nest site tending and defending



the embryos until hatching occurs (Andrews and Flickinger,  1973).



More than  one  female may spawn  in  the male's  nest,  and up to



12,000  embryos  have  been found  in  one nest  (Markus,  1934),



indicating  that several females contribute to the embryo mass.



     Egg counts in wild mature females have ranged from 800 to



9,000  eggs per female,   depending  on size  of female,  water



temperature, geographic  strain, etc.    Unpublished data  indicate



an  average of 258 eggs/spawn  and  an  average  of 3095  total



eggs/female during  100  days of spawning activity (Olson, 1974).



     For spawning  tanks,   a  standard tank is  divided  into 4



chambers as previously  described.   Each  spawning chamber is  then



approximately 15 cm x 30 cm.  A spawning substrate, and a pair of



sexually mature fish is placed in each section,  for a total of





                              22

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four pairs per spawning tank.  There are 24  spawning tanks in the



ERL - D system, for a total of 96 spawning pairs.



     Separation into spawning  pairs is not essential.   Other



investigators have reported success with male to female ratios of



2/4 (Olson,  1974), 3/6 (Benoit and  Carlson,  1977), and 4/10 -  15



(Mount, 1971).  The males are territorial,  so at least  as many



spawning substrates must be provided as there are males in the



tank to achieve optimum egg production.



     Paired spawning reduces fighting and competition  between



males.   When  ERL  - D began to  use  the  paired method,  the number



of embryos produced  in the culture unit  almost doubled  (Benoit,



1982).   Paired  spawning also allows the culturist to  follow the



fecundity of  individual  fish.   Sterile or  "spawned out" fish



identified through daily spawning records can then be  replaced to



maintain egg  production.  A good  procedure is to review daily



spawning records  every three weeks.  If a pair has not  spawned



during that period,  discard both male  and female and  replace



with new spawners.







4.4  Embryo  Incubation







     Each morning at approximately 10:30 am (7 days/week), all



spawning tiles are checked for the presence of embryos.  It is



not  necessary to remove the tile from the tank to  see the



embryos. Gently  running the fingertips on the undersurface of



the  spawning  substrates  will  reveal their presence.    If  no





                             23

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embryos are present, the spawning substrate is left in the tank.



This creates less disturbance than daily removal, inspection,  and



replacement of  each substrate  (See Photograph  #6).



     When checking for embryos,  estimate and record the number



deposited by each spawning pair. Provide fresh tiles  for each



chamber from which  embryos were  removed.  Researchers  requiring



embryos less than 24 hrs old for testing can take charge of  the



embryos at this point.   If the embryos are to  be  incubated, leave



them on the spawning  tiles, and place the tiles  in the dated



hatching tray.   Wipe the tops of the tiles clean  with  a sponge



to reduce   transfer of  sediment to the hatching pan.  Place tiles



on end, with two tiles pushed together to form a circle,   and



place  an  airstone between them to circulate   water.   (See



Photograph #7).   Inspect embryos  daily,  and remove any dead ones



with tweez€srs   to decrease the spread of fungus.   (Nonviable



embryos are opaque or  clear with a white "dot"  inside where  the



yolk has  precipitated.  Viable  embryos will be clear for  the



first 36-48 hrs until they reach  the eyed  stage).  If  fungus



has attacked more than 50% of the embryos on a  particular tile,



discard the entire spawn on that  tile.



     At 22°C   embryos will begin  to   hatch in 5 days.     If



embryos have been removed  from  spawning chambers each day, so



that the age of each group   is known to within 24 hours, then  95%



should hatch within a 24  hour period.  If a toxicity test is



planned that calls for less than 24  hr. old larvae, provide  the



fish   to  the researchers  at this point.  If larvae are to be





                             24

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Photograph 6.  Fathead minnow embryos on the  underside of a
spawning substrate.

                             25

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                                         f
                                         •S
                                         -5

                                         3
                                        S
                                        -5

                                        I

26

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reared to 30 days old for testing as juveniles, or to adulthood



for future brood  stock, remove them from the hatching tray using



a large bore, 50  ml volumetric pipette.  Count larvae into lots



of 250,  and stock into rearing  tanks  (15 gal  tanks with appx 20



cm water depth).  Label each rearing tank  with the date of  hatch.



Thirty day old juveniles that weigh approximately 150 mg each



(wet weight)   are the desired goal, although fish in the range of



100 to 300 mg per fish are acceptable  (EPA,  1982).



     Clean spawning tiles after  each  use  by soaking them in a



mild HTH13 bath (sodium hypochlorite,  appx 12 g HTH/liter) for 1



hour.  Follow with  a tap water  rinse,  and then soak tiles in



appx. 5 x 10~3 M sodium thiosulfate for at least 10 minutes to



remove residual chlorine.  After a final rinse in  culture  water,



set tiles aside to dry and store for reuse.








4.5  Larval Feeding







     All fathead minnow larvae less than 30 days old are fed live



brine shrimp  twice each day.  Since  larvae begin  feeding  during



the first day of hatch,  feeding must  start immediately. It is



important that the brine  shrimp  nauplii  be small  enough for the



fathead larvae to ingest  (Norberg and Mount, 1985).  Gape size of



larval fathead minnows  is  in the range of .24 -  .28 mm, so the



"width" of the nauplii when  offered  must  be slightly less than



this  for the nauplii to be ingested.    Growth rates depend on



the brine  shrimp nauplii used,  since different strains of the





                              27

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cysts  vary in nutritional  adequacy (ASTM Draft No.  2,  1984).



The brand now  in use by the ERL  -  D  fathead culture unit  is



Biomarine1  which has yielded more uniform growth than some



others.  Any strain that offers a good hatching percentage,  small



size of nauplii upon hatch,  nutritional adequacy, and freedom



from contaminants  will yield good results.  Some culturists also



supplement the larval diet with powdered or flake foods to insure



a more rounded diet.  Opinions vary as to the efficacy of this



supplementary feeding.  Some  trout  chow  formulations have proven



to be inadequate  fathead minnow  diets in experiments  in our



laboratories.



     Conditions for hatching each strain of cysts are usually



provided by the supplier.  Live nauplii are harvested from 24 -



48 hrs after set-up at 25°C. To harvest,  remove the air tube from



the hatching jars approximately 15 minutes  before feeding,  to



allow the live shrimp to settle.   Unhatched cysts will form a



brown layer at the  bottom of the  hatching jar, while the live



shrimp will form a bright orange  layer  just  above the unhatched



cysts.   The layer  of live  shrimp is  then siphoned  from the



hatching jars into a  1 liter beaker.  Rinsing the brine shrimp  is



not necessary  in flow-through systems with large tanks.   In



static  systems rinsing  is  desirable  to avoid salt  build up.



Exact quantification  of shrimp fed to each tank is not necessary



as long as all fish receive an adequate amount.  Inspect tanks  10



- 15 minutes after  feeding.  If all shrimp  have  been eaten  in



this short time, provide more.  It is  important that larvae  be





                             28

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feed ad libitum,  especially during the first few weeks of life.



Approximately 50 ml of the concentrate is fed  to each tank of



250 larvae, twice per day.   Larvae less than 1 week old require



slightly less.  Nine 1.5 liter jars started with approximately 25



cc of eggs  each,  provide two  feedings per day for 60 tanks of 250



larvae each.  Different brands,  feeding schedules, and stocking



densities  may alter the amount required.






Adult Feeding






     Feed spawning pairs,  replacement brood  stock,  and any other


                                            T ^
fish  over  30  days old  frozen brine shrimp-1-  twice  per day.



Allow frozen brine shrimp to thaw slightly  (not fully) for ease



of handling.  Feed fish   ad libitum, with each spawning pair



receiving 1/8 - 1/4 teaspoon,  and other tanks according to number



of fish. A rule of thumb is that the right amount of food will



be consumed in about 10 minutes.






4.7  Disease






     Discard any diseased lots of fish  and disinfect the tanks



with hypochlorite.  To minimize the risk  of spreading disease



between tanks, disinfect all nets, siphons, brushes, and other



tools  if disease is thought  to  be  present.  Adding  fifty ml



formalin16, and  10 ml Roccal II17  to  50 liters  of  water in a



plastic garbage can makes a handy disinfectant bath.





                              29

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     Fungus on eggs is not considered a disease (Mount,  1971).



Fungus is ubiquitous in the aquatic environment, and its presence



should not be considered a valid reason  to discard  all



associated  embryos unless the  labor involved in picking and



sorting out  the bad embryos is not worth the return at hatch.



     Examine fins, skin,  and gills  of fish from random tanks



monthly for parasites.  Perform other disease checks sufficient  to



assure that disease is not a problem.  If present, it is best  to



destroy fish from  infected tanks.  Treatment is in order only  if



the fish cannot be sacrificed.



     Treated  fish are  rarely,  if  ever,  used  in  bioassays.



Disease treatment does have a place,  however, for fish brought  in



from the wild.   When  bringing in wild brood stock the following



prophylactic treatments  may be useful.    First,  bring fish   to



25°C from  their arrival temperature,  at a rate not to exceed 2°C



per day.  After an extra  day at 25° for acclimation, disease



treatments  begin, following  the methods of ORSANCO (1974).



Perform disease treatments on incoming fish in a quarantined



area,  away  from the main culture  unit, if possible.



     Treat  fish  with potassium permanganate for external



parasites.  Using a 1% KMnO4 solution, add 1.0 ml per liter  of



water.   The water will turn dark purple,  and must  be aerated



vigorously  during  the  treatment.   After 1/2 hour, neutralize the



KMn04  with  .01 ml/liter of 0.1N sodium thiosulfate.  Within 15 -



20  minutes the solution will turn yellow - brown.   Siphon the



tank or drain it down  as far as  possible without stressing the





                             30

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fish,   and then  refill.  Give this treatment  for two consecutive



days,  and  curtail feedings during this time.



     Once  a  day for  the next four  consecutive days  use a



tetracycline  bath,  at the rate of 25 mg/liter,  for  bacterial



control.   After a one hour static  bath turn the water on, so that



the initial dose is being slowly diluted.  Do not drain the tanks



after this treatment,  and feed the fish  during the treatment.



Tetracycline  can also  be  added to the feed as an antibacterial



drug  against Aeromonas,  Hemophi].us,  and  Pseudomonas.   The



recommended  dose is 2.5 - 3.75 g/45 kg fish/day  for 10 days, in



feed (Schnick, et al, 1986).







4 .8  Gene pool







     The Environmental  Research Laboratory -  Duluth periodically



(every 2 years) mixes  existing brood stock  with healthy wild



minnows to  eliminate the risk of developing  a homogenous  strain



(Benoit,  1982). Homogenous  genetic stock may provide smaller



variance  in test results, but it correspondingly reduces the



strength  of possible  inferences that can be made.   Excessive



inbreeding leads to increased rate of genetic deformities.  When



bringing in fish for the infusion  of new genes make every attempt



to obtain the healthiest  fish possible.  Fish  that  show any sign



of disease, or that have  a known  history of exposure to disease,



dissolved  oxygen  stress, or exposure to other  environmental



perturbations must be  rejected.  Fish that are selected for the





                              31

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outbreeding process must be subjected to the previously described



temperature acclimation and disease treatment measures.



     After acclimation and treatment,  set up  the new brood stock



in an area separate from the main culture unit.  Maintain these



fish in quarantine, and breed through the second generation using



the same  procedures as in the main culture unit.  When  second



generation fish are sexually mature,  mix in spawning pairs with



original brood  stock, in combinations  of new male/existing female



and new female/existing male. The generation resulting from these



crossings  will  be the new brood stock for the main culture unit.



Replace one  fourth of the spawning pairs  in the culture unit



every  three  weeks until all  pairs have  been changed. Consult



researchers  using the fish in toxicity tests so that they are



aware that new genes will be entering the pool.







4.9  Record keeping







     Maintain  records on the  estimated number of eggs spawned



each day by each spawning  pair.  Also record  the  date that each



spawning pair was introduced,  and the dates on which each holding



tank was  stocked with larvae.  Measure and record   routine water



chemistries (pH and D.O.)  at least twice weekly on random  tanks.



Records on source  of new brood stock, date of arrival, disease



treatments,  and procedures  used for mixing into the gene pool are



also important. Record all mortalities in the culture unit, and



observe fish  daily for abnormal appearance or  behavior.





                              32

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                       LITERATURE CITED
Allison,  D.T., and R.O.  Hermanutz.  1977.   Toxicity of diazinon to
brook trout and fathead minnows.  U.S. Environmental Protection
Agency.  EPA - 600/3  - 77 - 060.


American Society for Testing and Materials (ASTM). 1984. Proposed
new standard practice for the use of brine shrimp nauplii as food
for test animals in  aquatic toxicology. Draft No.  2, August, ASTM
Subcommittee E47.01  on Aquatic Toxicology.

Andrews, A.K., and S.A. Flickinger. 1973. Spawning  requirements
and characteristics  of  the fathead minnow.  Proc.  27th. Ann.  Conf.
SE Assoc.  Game and Fish Comm. pp 759 - 766.

Benoit, D.A.,  and R.W.Carlson. 1977. Spawning success of fathead
minnows on selected artificial  substrates.  Prog. Fish.  Cult.
39(2).67 -69.

Benoit,  D.A.  1982. User's guide for conducting  life  cycle chronic
toxicity tests with fathead  minnows  (Pimephales promelas).  U.S.
Environmental Protection Agency. EPA - 600/8 - 81 -  Oil. 17pp.

Brungs, W.A. 1971a.  Chronic effects of elevated temperature on
the fathead minnow  (Pimephales promelas  Rafinesque). Trans. Am.
Fish.  SOC.  100(4). 659 - 664.

Brungs,  W.A.  1971b. Chronic  effects of low dissolved  oxygen
concentrations on the fathead minnow,  (Pimephales  promelas). J.
Fish.  Res.  Bd. Canada. 28. 1119 - 1123.

Drummond,  R.A., and  W.F. Dawson. 1970. An inexpensive method for
simulating diel patterns of lighting in the laboratory. Trans.
Am. Fish.  Soc. 99. 434 - 435.

Eddy,  S.,  and J.C. Underbill. 1974. Northern fishes. Univ. Minn.
Press,  Mpls.,  MN.  414pp.

EPA,  1982.  Culture of 30 day old fathead minnows  for use in acute
tests at ERL - D. Standard operating procedure #CR - 01 - 01.
U.S.  Environmental Protection Agency, Duluth, MN  55804. 2pp.

Flickinger,  S.A.  1969.  Determination of sexes  in the  fathead
minnow.  Trans. Am. Fish. Soc. 3. 526 - 527.

Gale,  W.F.,  and G.L.Bunyak.  1982.  Fecundity and spawning
frequency  of  the fathead minnow - a fractional spawner. Trans.
Am. Fish.  Soc. 111.  35 - 40.
                              33

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Glass, G.E.  1977. Identification and distribution of inorganic
components in water:  what  to measure?  Ann.  NY Acad. Sci.  298. 31
-46.

Hasler,  A.D.,  H.P.  Thomsen,  and J.C.Neess.  1946. Facts and
comments on raising two common bait minnows. Wis. Cons.  Bull.
#210  - A - 46.  12 pp.

Isaak,  D.  1961.  The  ecological life  history of  the  fathead
minnow,  P!roejoha].es P_rome_las  (Raf inesque). Ph.D. Thesis,  Univ.
Minn.  150 pp.

McCormick,  J.H., and R.F. Syrett. 1970. A controlled temperature
apparatus   for   fish  egg  incubation  and  fry  rearing.
manuscript.U.S. Environmental Protection  Agency, Duluth,  MN

McMillan, V. 1972. Mating of the fathead. Nat. Hist.  81(5).73 -
78.

Mount, D.I.  1971.  Tentative plans for the design and of operation
of a  fathead  minnow stock culture unit.  U.S.  Environmental
Protection Agency, Duluth,  MN  55804. 7pp.

Norberg,  T.J.,  and  D.I.  Mount. 1985. A new subchronic fathead
minnow (PiraepJiaJ^s P_rome,lasJ toxicity test.  Env. Tox. Chem. 4.
711 -718.

Olson, D.T.  1974.  Fathead minnow egg production as a function of
the number of females per spawning chamber.  Interim report of the
fathead  minnow methods study, Part I.  15pp. U.S. Environmental
Protection Agency, Duluth,  MN  55804.

ORSANCO.  1974. 24 hour bioassay.  Ohio River Valley Sanitary
Commission.  Cincinnati, OH 45202.  21pp.

Radcliff,  L.  1931.  Propagation of  minnows. Trans. Amer.  Fish.
Soc.  61.  131 -  137.

Schnick, R.A., F.P. Meyer,  and D.F.  Walsh. 1986.  Status of
fishery chemicals in 1985.  Prog. Fish. Cult. 48(1). l - 17.

Syrett,  R.F., and W.F. Dawson.  1972.  An inexpensive electronic
relay  for precise water temperature control. Prog. Fish.  Cult.
34(4). 241 - 242.

Syrett,  R.F., and  W.F. Dawson.  1975. An inexpensive solid  state
temperature controller. Prog.  Fish.  Cult. 37(3). 171 - 172.
                              34

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                      ADDITIONAL READING

Adelman,  I.R.,  and L.L.  Smith,  Jr. 1976.  Fathead minnows
(Piro^Ekf*.!6.^  E££3l®.ll:fL)  and  goldfish (Carass_ius auratus) as
standard  fish  in  bioassays  and their  reaction  to  potential
reference toxicants. J. Fish. Res. Board Can.  33.  209-214.

Andreasen,  J.K.  1975.  Occurrence  of the  fathead minnow,
Pimephales promelas,  in Oregon.  Calif.  Fish and Game. 61(3).  155-
156.

Andrews,  A.K.  1970.  The distribution and life  history of the
fathead  minnow (Pimejohaleis promelas Rafinesque) in Colorado.
Ph.D. Thesis.  Colorado State Univ. Fort Collins.  131pp.

Andrews,  A.K.  1970.  Squamation chronology of the  fathead minnow,
Pimephales promelas. Trans. Am. Fish. Soc.  99(2).  429-432.

Andrews, A.K. 1971. Altitudinal range extension  for the fathead
minnow (Pimephales promelas). Copeia. 1. 169.

Andrews,  A.K.  1972. Survival and  mark retention of  a  small
cyprinid  marked with flourescent pigments.  Trans. Am.  Fish.  Soc.
101(1).  128-133.

Bernstein,  J.W.,  and R.J.F.  Smith.  1983. Alarm  substance  cells in
fathead minnows do not affect the  feeding preference of rainbow
trout. Environ. Biol. Fish.  9(3-4). 307-311.

Brauhn, J.L.,  and R.A. Schoettger. 1975. Acquisition and culture
of  research  fish:  rainbow trout,  fathead  minnows,  channel
catfish,  and bluegills. US Env.  Prot. Agcy.  EPA-660/3-75-011.
45pp.

Brown, B.E.  1970. Exponential decrease in a population  of fathead
minnows.  Trans. Am.  Fish. Soc. 99(4).  807-809.

Chiasson, A.G., and J.H. Gee. 1983.  Swim bladder gas composition
and control of buoyancy by  fathead minnows (Pimephales  promelas)
during exposure to  hypoxia.  Can. J. Zool. 61(10).  pp 2213-2218.

Coble, D.W.  1970. Vulnerability  of  fathead minnows infected  with
yellow grub to largemouth  bass predation.  J.  Parasit. 56(2).  395-
396.

Coyle, E.E. 1930. The algal food of PiinejDha3.es promejjas. Ohio J.
Sci. 30(1). 23-35.

Dixon, R.D. 1971. Predation of mosquito  larvae by the fathead
minnow,  Pimephales  promelas Rafinesque. Manit.  Entomol.  5.  68-70.
                              35

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Flickinger,  S.A. 1973. Investigation of pond spawning methods for
fathead minnows.  Proc. 26th. Ann. Conf.  SE Assoc.  Game and Fish
Comm. 376-391.

Cast, M.H., and W.A. Brungs.  1973. A procedure for separating
eggs of the fathead minnow. Prog. Fish Cult. 35. 54.

Gee, J.H.  1977. Effects of size  of fish, water temperature, and
water  velocity  on  buoyancy alteration  by  fathead  minnows,
Pimephales promelas.  Comp. Biochem.  Physiol. A56(4). 503-508.

Gee, J.H., R.F.  Tallman, and H.J. Smart.  1978. Reactions of some
great plains  fishes to progressive  hypoxia.  Can. J.  Zool. 56.
1962-1966.

Grabar,  F.,  and R.A.  Flickinger.  1977.  Replicon number and growth
rate in fathead minnow  cells  cultured at 14 and 34°C. Cell Tissue
Kinet.  10(5). 505-507.

Gravell, M., and R.G. Malsberger. 1965. A permanent (epithelial)
cell  line   from   the   fathead  minnow    (Pi.m ep_ha 3. es
promelas)(cultivation method, temperature control, chromosome
analysis, virus susceptibility). Ann NY Acad Sci. 126(1). 555-
565.

Guest,  W.C.  1977.  Technique  for collecting  and incubating eggs of
the fathead minnow. Prog. Fish. Cult.  39(4). 188.

Hart, J.S.  1947. Lethal temperature relations  of certain fish of
the Toronto region. Trans. Roy. Soc. Can. 91(3). 57-71.

Held, J.W. 1971. Some ecological aspects of the fathead minnow,
Piro^Eli^i6-3- PJ-!P.S!®.l3ts Rafinesque,  in  North Dakota saline lakes.
Ph.D. Thesis. North Dakota State Univ. Fargo, ND. 80pp.

Held, J.W.,  and J.J. Peterka. 1974. Age,  growth,  and food habits
of  the  fathead minnow,  P_imej3haJ.es  p_romeJLa£, in North Dakota
saline lakes. Trans. Am.  Fish. Soc. 103(4). 743-756.

Hendrickson,  G.L.   1979.  Ornj-thodi-p^ostomum ptychocheilus;
migration to the brain of the fish intermediate host, Pimephales
promelas.  Exp.  Parasit. 48. 245-258.

Hoffman,   G.L.   1958.   Studies   on  the  life  eye], e  of
2£Hi^ho^i£l.2st°lH51  ptychocheilus (Faust)  (TrematodarStrigeoidea)
and the "self cure"  of  infected  fish.  J.  Parasit. 44(4).  416-421.

Ingram, R., and W.D. Wares,II.  1979. Oxygen consumption in the
fathead minnow (Pimep_hales p_romelas Rafinesque)  II: Effects of
pH, osmotic pressure,  and light level.  Comp. Biochem.  Physiol.
62A. 895-897.
                              36

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Kelly, R.K., B.W. Souter, and H.R. Miller. 1978. Fish cell lines:
comparisons of CHSE-214, FHM, and RTG-2 in assaying IHN and IPN
viruses.  J. Fish. Res. Board Can.  35(7).  1009-1011.

Klak, G.E. 194Q.Neascus infestation of blackhead,  blunt nosed,
and other forage minnows.  Trans. Amer. Fish. Soc.  69.  273-278.

Klinger,  S.A., J.J.  Magnuson, and G.W. Gallepp.  1982.  Survival
mechanisms of the central  mudminnow  (Umbra  limi), fathead minnow
(Pimephales promelas),  and  brook stickleback  (Culea  inconstans)
for low oxygen in winter. Environ. Biol. Fish. 7(2). 113-120.

Konefes,  J.L., and R.W. Bachmann.  1972.  Growth of the fathead
minnow  (Pimephales promelas) in tertiary treatment ponds.  Proc.
Iowa Acad. Sci.  77. 104-111.

Lemke,  A.E.,   W.A. Brungs,  and B.J.  Halligan. 1978. Manual for
construction  and operation of toxicity testing proportional
diluters. US Env. Prot. Agcy. EPA-600/3-78-072.

Lewis,  D.H.,  and N.L.  Savage. 1972.  Detection of antibodies to
Aeromonas iiquefjici.ens  in fish by  an  indirect flourescent
antibody technique. J. Fish. Res.  Board Can.  29(2). 211-212.

Lord, R.F., Jr. 1927. Notes on the use of the blackhead minnow,
PimephjQes P_romel.asj,  as a forage fish. Trans. Am. Fish. Soc. 57.
92-99.

Luoma,  M.E.,  and J.H.  Gee.  1980.  Seasonal factors affecting
buoyancy attained in still  water and current by fathead minnows,
Pimephales promelas. Can. J. Fish.  Aquat.  Sci.  37. 670-678.

Manner,  H.W.,  and C.M. Casimira. 1974. Early embryology of the
fathead minnow Pimephales promelas Rafinesque. Anat. Rec. 180(1).
99-109.

Manner,  H.W.,  and C. Muehlman. 1975. Permeability and uptake of
3H-uridine during teleost embryogenesis. Sci. Biol. J.  1(3). 81-
82.

Manner,  H.W.,  M. VanCura,  and C.  Muehlman.  1977.  The
ultrastructure of the  chorion of  the fathead minnow,  Pimephales
promelas. Trans.  Amer. Fish. Soc.  106(1).  110-114.

Marcus,  L.F.,  and J.H. Vandermeer.  1966.  Regional trends in
geographic variation. Syst.  Zool.  15(1).  1-13.

Markus,  H.C.  1931.  Life history  of the blackhead minnow,
Pimephales promelas. Copeia. 3. 116-122.

Mayer,  F.L. 1976. 2,4-D reduces SajDro_legji.ia on fathead minnow
eggs. Prog. Fish. Cult.  38(1). 19.

                              37

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Merritt,  R.B.   1972.  Geographic  distribution  and enzymatic
properties of lactate  dehydrogenase allozymes  in  the  fathead
minnow,  Pimephales promelas. Am. Nat. 106(948).  173-184.

Meyer, F.P. 1958. Helminths of fishes from Trumbull Lake,  Clay
County,  Iowa. Proc. Iowa Acad. Sci. 65.  477-516.

Minchew, C.D. 1981. Unicauda magna sp.n. (Protozoa:  Myxozoa):  A
new  Myxozoan from  the  fin  tissues  of the  fathead  minnow,
Pimephales promelas Rafinesque. J. Fish.  Dis.  4(6). 513-518.

Ming, F.W,  and D.L.G. Noakes.  1984. Spawning  site selection and
competition  in  minnows  (P_imepjia_les notatus  and P^ prgmejlas)
(Pisces, Cyprinidae). Biol. Behav.  9(3).  227-234.

Mitchell, A.J., C.E. Smith, and G.L. Hoffman. 1982. Pathogenicity
and histopathology of  an unusually  intense  infection of white
grubs  (Posthodi.p_lostomum m. m_in_imum)   in the fathead  minnow
(Pimephales promelas).  J.  Wildl. Dis. 18(1).  51-57.

Mitchell, L.G.,  C.L. Seymour,  and J.M.  Gamble.  1985. Light and
electron microscopy  of Myxobolus hendri.cksoni (Myxozoa,
Myxobolidae)  infecting  the brain of  the fathead minnow,
Pimephales promelas Rafinesque. J. Fish.  Dis.  8(1). 75-89.

McCarraher,  D.B.,  and  R.   Thomas.  1968.  Some  ecological
observations  on  the fathead minnow,  Pimephales promelas, in the
alkaline waters of Nebraska. Trans. Am.  Fish.  Soc.  97(1).  52-55.

McMillan,  V.E.  1972.   An  ethological  study of agonistic and
reproductive  behaviour in the fathead minnow (Pimephales promelas
Rafinesque).  Ph.D. Thesis.  Univ. Saskatchewan, Saskatoon, Sask.,
Canada.  147 pp.

Nagel, T. 1976.  Technique for collecting newly hatched  fathead
minnow fry. Prog. Fish.  Cult. 38(3).  137.

Niazi, A.D.   1964. The  development of  the Weberian system and
early embryology  of  Pimeghal.es P_rome_las  (Osteichthyes:
Cyprinidae).  Diss. Abstr.  25(4). 2674.

Peterson, D.L.,  and F.A. Hennagir.  1980.  Minnesota live  bait
industry assessment study.  Proj.  #30261-R.  Minn.  Dept. Natl.
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Prather, E.E. 1958. Preliminary experiments on  winter  feeding
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                             38

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Radabaugh/  D.C. 1980. Changes  in minnow,  P^me^
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Sykora, J.L., E.J.  Smith,  and M. Synak. 1972. An inexpensive fish
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