Plankton Ecology

66
Food chains and webs - simple

Organisms can be organized into

food chains & food webs of
varying complexity
 67
Food webs - sublime

• Probably more realistic

ecologically, but perhaps
useless to managers
• But be aware of not-so-
obvious complexities

68
Food and energy transfers in lakes

Microbial
loop
 69
Algal primary productivity

Photosynthesis
-Light - Temperature
-Nutrient - Chronic toxicity
-Velocity

Washout
Loading Turbulent -Velocity
diffusion
Algal biomass -Available substrate

Mortality Sinking
Grazing -Acute toxicity - Velocity
-High temperature - Stress
Respiration/Excretion

70
Population models
How do populations increase?
Birth, immigration
How do populations decrease?
Death, emigration

r=b-d
(assumes a closed population)

r = rate of change in N
b = proportional birth rate
d = proportional death rate
 71

12000

10000
Population Size (N)

8000 r=0
b=d
6000
Good times Bad times

4000 r>0 r<0

b>d b<d
2000

0
0 5 10 15 20 25 30 35
Time

72

changes in populations
are multiplicative
∆N/∆t = dN/dt = rN
a large population can add
tremendous numbers of individuals
in a short period of time
 73

r = rate of increase
instantaneous rate of change of
N with respect to time (slope tangent
to N vs. t graph)

r = ln Nt – ln No change
in N

t change in time

N0 = population at time 0
Nt = population at time t

Temperate-Lake Phytoplankton: 74
A Study System in Ecology

• Abundance varies >= 1000X and composition

undergoes predictable seasonal shifts
• Observations of patterns led to hypotheses about
why different populations increased and crashed at
different times...
• Rapid life cycles, simplicity of culture allowed for
experiments to determine processes underlying the
patterns
 75
Competition, in part, structures algal assemblages.

76
Exploitative competition — competition through
removal of resources

Interference competition — competition through

direct interference

Allelopathy – the direct effect of competitors through

the release of chemicals that inhibit other species
 77
How do you know who wins in competition?

Early experiments (prior to 1970s) used circular logic.

The superior competitor is the one that will

displace all other species

Put two species together on a limiting resource, let it

run for many generations (usually used aquatic
protists, so several generations wasn’t someone’s
lifetime).

The species that displaced all other species was

declared to be the superior competitor

78
Is there a way to tell who will win before you put
them together?

Tilman’s Mechanistic Theory of Resource Competition

Tilman 1982 — several papers prior to that

assessed competitive ability of algae based on
their physiology.
 79
Tilman tested his theory using diatoms from
Lake Michigan
Asterionella and Cyclotella
Grew each alone in the chemostat under P-limited
conditions

Diatoms are Phosphate limited 80

No matter where
you draw the
mortality curve,
Asterionella
µ

should always
out compete
Cyclotella under
conditions of P-
limitation
PO4 (µmol/L)

He ran the experiment 9 times with Asterionella and Cyclotella

together under P-limited conditions and Asterionella always
displaced Cyclotella
Diatoms can also be limited by Si. 81

No matter where
you draw the
mortality curve,
Cyclotella should

µ
always outcompete
Asterionella under
conditions of Si
limitation.

SiO2 (µmol/L)

Asterionella is a good competitor for P, but relatively poor for Si

Ran this experiment 6 times—Cyclotella won 5 out of 6 times.

82
Asterionella is a better competitor for P

Cyclotella is a better competitor for Si

Who wins depends on ratio of two

nutrients.
 83
Tilman’s experiments were stimulated in part by
two papers that came out in the early 1960s.

Competitive exclusions principle (Hardin 1960)

The number of species coexisting at equilibrium
will be limited to the number of simultaneously
limiting resources.

For phytoplankton, that is typically 3-4.

84
Paradox of the Plankton (Hutchinson 1961)
If there are only 3-4 limiting nutrients, how can
there be dozen or hundreds of phytoplankton
species present in a lake?

Answer: conditions change faster than any one

species can outcompete the others.

Non-equilibrium conditions
 85
Tilman and others also looked at N:P ratio, Light:P etc.

For example bluegreens tend to out-compete other species

at low N:P ratios. They can fix atmospheric nitrogen.

RECAP: What
drives seasonal
patterns of
abundance and
distribution??

Abiotic Factors
Competition
Grazing

86
Other regulators of lake productivity - Grazing

Top-down Model
Bottom-up Model
• High rates of nutrient driven algal
growth is removed by intense • Nutrient inputs
zooplankton grazing pressure (usually
cladoceran Daphnids) drive algal growth
> Fishless lakes with low zooplankton predation • Classic Pyramid
> Lakes where planktivorous fish are regulated
by predatory fish (game fish) –usually by
intensive control
> In these cases, algae are not nutrient limited

N+P
 87
Potential Top-down effects on food chains
Low HIGH
Predators Predators

HIGH Low
Planktivores Planktivores

Low HIGH
Zoops
= Smaller Zoops Larger Zoops
=
less grazing more grazing Zoops

HIGH Low Secchi Higher Secchi Low

Algae O2 stress less O2 stress Algae
high pH ?? lower pH ??

88
Grazing
Rotifers, copepods and cladocera all include species that
consume algae
http://www.eeb.cornell.edu/hairston/hairston.html
www.uv.es/~ciros/zoopl_en.html http://dnr.metrokc.gov/wlr/waterres/lakes/daphnia.jpg

Brachionus (300 µm) Daphnia

Diaptomus (< 2 mm)
(2-3 mm)
Cladocera, especially Daphnia, are the main grazers in
aquatic systems.
 89
Strong correlation between peak abundance of
Daphnia and timing of annual Clearwater phase.

But, correlation is not causation.

In early 1980s, much debate about what caused Clearwater

phase, was it abiotic, was it nutrient limitation, was it grazing?

90
Indirect evidence for role of grazing:

Primary productivity per unit chl-a is highest in the summer

This suggests that the cells that

are there are in good condition

The ratio of pheopigments (degradation product of Chl-a)

to Chl-a is highest in the summer

Guts of zooplankton are acidic, acid

degrades Chl-a; lots of pheopigments
suggests that algae are being eaten
 91
Can also measure grazing directly:
Tessier et al. (2001)—Limnology and Oceanography
http://www.cladocera.uoguelph.ca/taxonomy/daphnia/default.htm

www.microscopy-uk.org.uk/ mag/imgjun99/cerio.jpg

Ceriodaphnia Daphnia Daphnia Daphnia

reticulata ambigua mendotae pulicaria
(< 0.8 mm) (1.4 mm) (1.9 mm) (2.5 mm)

Cultured each species of grazer in the lab, and then added

them into enclosures in the lake

Duck Lake, MI 92

Pump water from the lake, through a mesh,

into the bags

Add the grazers to the bags

Sample bags
every 3-7 Replicates of each treatment
days for 40
days
 93
What was the impact
of the grazers on the
algae?

94
More than 50 different types of phytoplankton were observed
in the experiment

Small chrysophytes, chlorophytes, dinoflagellates and

centric diatoms were quickly eliminated from all treatments

Other taxa such as Dinobryon and Oocystis increased in all

treatments

Many algae showed different responses in the treatments

with the small-bodied grazers vs. the large-bodied grazers
 95
Take-home message is that the Tessier et al. and many
other studies have shown:

Lots of Grazing on:

Single celled green algae Small Diatoms Dinoflagellates
microbes.limnology.wisc.edu/outreach/ images/peridinium.gif
http://www.ucmp.berkeley.edu/greenalgae/chlamydomonas.jpg

Not as much grazing on:

Cyanobacteria Gelatinous greens
http://www.biol.tsukuba.ac.jp/~inouye/ino/g/chl/oocystis.jpg

BUT, the loss rate for the algae often depends on the grazer

96
Trade-offs again, some species that are good competitors for
limiting nutrients, are also vulnerable to being eaten.

Why are some algae less vulnerable to grazers than others?

1. Difficult to handle

2. Bad taste or toxic

3. Nutritionally poor food

 97
1. Handling time: Infante and Abella (1985)—Limnology and Oceanography

Studied Daphnia feeding on Oscillatoria from Lake Washington

http://www.cladocera.uoguelph.ca/taxonomy/daphnia/default.htm www.ucmp.berkeley.edu/bacteria/ oscillatoria2.jpg

Daphnia pulicaria Daphnia thorata Oscillatoria

As number of filaments/ml increased from 0-400,

growth rate of both Daphnia species declined.

Both Daphnia species spent a lot of time rejecting the algae

98
2. Some algae are toxic: Lampert (1987)
New Zealand Journal of Marine and Freshwater Research

100
www-cyanosite.bio.purdue.edu/ images/lgimages/microcy3.jpg

80
Microcystis
% survival

60 No food

40

20

0
0 1 2 3 4 5
TIm e (days)

Microcyctis produces a toxin

But toxicity depends on both the algae and the grazer 99

Ferrão-Filho et al. 2000

Freshwater Biology

3 strains of
Microcystis

4 species of
cladocera

100
3. Differences in nutritional quality:

Essential fatty acids are needed for membranes and hormones.

Algae differ in their content, Diatoms have many fatty acids,

blue-greens have little.

Fatty Acids Cyanobacteria Greens Diatoms

Linoleic yes yes yes
Eicosapentaenoic no yes yes
Docosahexaenoic no no yes
 Is there any benefit to having grazers around? 101

Lehman (1980) —

Found microzones of N and P around Daphnia

Epilimnion zooplankton supply 10 time more P and 3 times
more N to the epilimnion than all other sources combined

Porter—several papers in the 1970s and 80s

Fed Chlamydomonas to Daphnia

Suggested that the algal cells could pass through the gut
without being assimilated.
Get nutrients from the gut.
This work was pretty much forgotten about until recently

102
In summary, seasonal succession is controlled by:

Abiotic factors Competition Grazing

Importance of each factor varies with time of year.

 103
Predation: Predators of zooplankton in Lakes and Ponds
Vertebrates : www.cnr.vt.edu/efish/families/ bluegill.html

http://www.potomacriver.org/images/biology/Leptodorabig.jpg
Invertebrates dnr.metrokc.gov/wlr/waterres/ lakes/biolake.htm

www.zi.biologie.uni-muenchen.de/ .../feinstruktur/em.htm

Find prey by vibration or

chemical cues
www.fs.fed.us/wcnf/fishing/ lakes/lake07.htm

Use vision to find prey

Tend to select smaller
Tend to select larger prey
prey

104
Predatory Invertebrates Insects
Notonecta--
backswimmer www.biol.lu.se/funkmorf/ vision/dan/prey.html

Dyticid
beetle larva

http://www.fhsu.edu/biology/thomasson/AquaticInsects/HerlOdonata/Herl.htm

Dragonflies
www.zi.biologie.uni-muenchen.de/ .../feinstruktur/em.htm

Chaoborus —
phantom midge
 105
Predatory Invertebrates Rotifers

Asplanchna Synchaeta

http://www.microscopy-uk.org.uk/mag/wimsmall/extra/rotif2.html

106
Predatory Invertebrates Crustaceans--Cladocera
http://www.potomacriver.org/images/biology/Leptodorabig.jpg

Leptodora Polyphemus

Cercopagis

Bythotrephes
 107
Predatory Invertebrates Other Crustaceans
Epischura

www.potomacriver.org/ ZoopIndicators2.htm
Mesocyclops

www.venturenorth.com/tlca/ pictures/mysis.jpg

Mysis

108
Predatory Invertebrates Other groups
ebiomedia.com/gall/classics/ Plan/planaria.html

Flatworms

www.microscopy-uk.org.uk/ mag/artdec99/mite3.html

Water mites
Planktivorous fish select large prey items 109

110
Size-Selective Predation
First discussed by Hrbáček (1962)
Rozpravy Ceskoslovenské Akademie Ved, Rada Matematickych a Prirodnich Ved

In ponds with < 700 fish/ha (minnows and perch), dominance

by large zooplankton:
Daphnia pulicaria (2.0-2.3 mm)
Daphnia longispina (1.4-1.8 mm)

In ponds with > 30,000 fish/ha dominance by small

zooplankton:
Bosmina longirostris (0.3-0.4 mm)
Daphnia cucullata (0.7-0.8 mm)
 111
Why do you find species A, B, and C in one pond
and literally meters away find species X, Y, and Z?

34 species of Daphnia in North America, but only get 2 or

3 per lake. Maybe as many as 5 or 6.

Often seasonal succession of zooplankton within a lake,

but we will focus on differences among lakes

As with the phytoplankton, population dynamics and

community structure are influenced by abiotic factors,
predation and competition.

112
Pattern Observed by Brooks and Dodson (1965)
Surveyed many lakes in New England and found that
Daphnia, Epichura and Mesocyclops were absent from lakes
containing Alosa pseudoharengus
www.potomacriver.org/ ZoopIndicators2.htm

Lakes with alewife were dominated by

Bosmina, Ceriodaphnia and Tropocyclops
www.potomacriver.org/ ZoopIndicators2.htm
 Alosa pseudoharengus 113
www.anr.state.vt.us/dec/waterq/ ans/ans-index.htm

Anadromous marine fish that feeds and grows to

maturity in the ocean, then migrates into freshwater to
spawn. But some populations have become “landlocked”
and live their whole life in lakes.

Unlike other planktivores like bluegill, Alosa will cross

the thermal barrier to feed

114
Crystal Lake, Connecticut

Brooks sampled the plankton in 1942

Alosa aestivalis invaded the lake in the 1950s

Dodson was an undergrad at Yale when the

lake was sampled in 1964
 Brooks and
115
Dodson 1965
(over 1350 citations)

To explain their
results they
proposed size-
selective
predation and the
Size-Efficiency
Hypothesis

Galbraith (1967) 116

Looked at feeding by rainbow
trout and yellow perch in Stager Lake

In Lake
In Guts
Frequency (%)

(Daphnia)
 117
Size-Efficiency Hypothesis
Brooks and Dodson 1965

1) Planktonic herbivores all compete for the

fine particulate matter of the open water

2) Larger zooplankton do so more efficiently and can

also take larger particles

3) Therefore, when predation is of low intensity the small

planktonic herbivores will be competitively eliminated by
large forms (dominance of large Cladocera and calanoid
copepods).

118
Size-Efficiency Hypothesis (continued)
Brooks and Dodson 1965

4) But when predation is intense, size-dependant predation

will eliminate the large forms, allowing smaller zooplankton
to dominate.

5) When predation is of moderate intensity, it will fall

more heavily upon the larger species and keep their
populations low, so that the smaller species are not
eliminated.
 119
What have we learned since 1965?

Pattern vs. Process

1) Planktonic herbivores all compete for the fine

particulate matter of the open water

Only when resources are limiting

2) Larger zooplankton do so more efficiently…

Not necessarily

2) …and can also take larger particles

But large zooplankton may miss the smaller particles

120
3) Therefore, when predation is of low intensity the small
planktonic herbivores will be competitively eliminated by
large forms (dominance of large Cladocera and calanoid
copepods).

Competitive dominance is not only a function of

body size

When predation is of low intensity on the large

grazers, it is also of low intensity on big
predators like Leptodora, Mesocyclops and
Epichura

Leptodora, Mesocyclops and Epichura eat small

planktonic herbivores
 121
4) But when predation is intense, size-dependant
predation will eliminate the large forms, allowing
smaller zooplankton to dominate.

Eliminating the larger forms includes eliminating

the predators of the smaller forms

5) When predation is of moderate intensity, it will

fall more heavily upon the larger species and keep
their populations low, so that the smaller species
are not eliminated.

To fully explain size structure of the zooplankton

assemblage, need to consider basin shape,
zooplankton behavior, competition and predation
(including type of predator)

122

There have been dozens if not hundreds of experiments

designed to investigate competitive dominance among
different zooplankton taxa.

Many studies have focused on large cladocerans (e.g.,

D. pulicaria, D. dentifera) because they are grazers
capable of reducing phytoplankton down to limiting
levels. If edible algae is abundant, no competition.

How do you measure competition among zooplankton?

 123
Either in the lab or in the field, set up replicate
enclosures of the following treatments:

Treatment 1 2 3 4 5
Species 1 0 0 ½ 1 ½

Species 2 ½ 1 ½ 0 0

Total ½ 1 1 1 ½

Intra-specific Competition—competition within one species

Inter-specific Competition—competition between more than one species

Dodson (1972, 1974) 124

S E H from Brooks and Dodson

(1965)

http://campus.murraystate.edu/academic/faculty/Howard.Whiteman/whiteman.htm

“Mexican Cut”
alpine ponds
Rocky Mountain
Biology Lab

6 ponds
 125
Some ponds were dominated by ‘small’ species
Daphnia minnehaha or Daphnia rosea

Some ponds had the “large” species Daphnia

middendorffiana but not the smaller species
http://www.cladocera.uoguelph.ca/taxonomy/daphnia/default.htm

126
Used “cage” experiments to transplant the
Daphnia among ponds

D. rosea pond

D. middendorffiana pond

the smaller species had no problem living with

the larger species in either pond.
 127
But, when he added Diaptomus shoshone to the cages…

D. rosea pond

D. middendorffiana pond

…the smaller species was excluded within one month

128
What controls the distribution of Diaptomus
shoshone?

http://campus.murraystate.edu/academic/faculty/Howard.Whiteman/research.html

Predation by vertebrates
 129
Bengtsson 1987 surveyed the literature and
collected data for the outcome of 20
competition experiments among cladocera and
found:

Predictions of Tilman’s R* were supported in 68% of cases

Size-Efficiency Hypothesis was supported in 60% of cases

76% of experiments that used the same species, but had

different conditions (field/lab, spring/fall) had different
outcomes.

130
Competition between rotifers and cladocera:
Big Daphnia tend to outcompete rotifers.
Work done by J. Gilbert and students

Keratella
r = 0.46 / day without Daphnia
r = -0.11 / day with Daphnia

Questioned if this was exploitative

competition or interference competition
http://www.cladocera.uoguelph.ca/taxonomy/daphnia/default.htm

Remember:
Exploitative—removal of resources
Interference—direct interference
 131
Here “interference” means the rotifers get caught in
the filtering combs of the Daphnia and are either
eaten or damaged.

133
Why are fish size-selective?

1. Encounter frequency: Encounter of large prey is higher

than small prey

Reaction distance —how

close to the fish does a prey
item have to be for the fish
to see it and react to (eat)
it?
www.state.ia.us/.../fwb/fish/ iafish/sunfish/pkse.gif

Pumpkinseed Lepomis gibbosus

Confer and Blades 1975 (L&O)

 134
Reaction distance translates to overall volume searched,
which influences vulnerability of the prey

Reaction distance = radius

of sphere

Longer radius = higher

encounter rate

Visual field is roughly spherical with ~ 20º

posterior blind segment
Volume searched = volume of sphere = 4/3 π r3

135
2. Preferred prey —take the prey that provides the
greatest energy return for cost of capture/handing.
Werner and Hall (1974) Ecology

With abundant prey, biggest possible is better

80
Expected from
encounter rates
60
Observed in Diet
% in diet

40

20

0
Small (< 1.5 mm) Medium Large (>2.5 mm)

Fed fish choice of three sizes of Daphnia magna

Get exception to this “bigger is better” rule with small fish 136
because of gape limitation Li et al. 1989 Ecology
Walton et al. 1992 Ecology

+1 Fish > 76 mm Selects for

particular
animals

Selectivity
0 No choice
Index

Fish = 11 mm

-1 Avoids
Daphnia size (mm) particular
animals

Bluegills feeding on Daphnia Small fish don’t eat large prey

137
One consequence of size-selective predation is keystone
predation

Keystone Predation – predator preys on competitively

dominant herbivore, thereby increasing death rates and
facilitating persistence of the weaker competitor

Although zooplankton body-size is not a perfect

predictor of competitive ability, often the big species is
the dominant competitor.

Explanation for #5 prediction of the SEH (when

predation is of moderate intensity…smaller forms are
not eliminated)
 138
Most predatory invertebrates that feed in the
plankton tend to select prey that are smaller than
they are

So they select for small

species (e.g., Bosmina,
Ceriodaphnia, Rotifers)

Or juveniles of larger http://www.microscopyu.com/moviegallery/pondscum/crustacean/nauplius/

species (e.g., nauplii,

juvenile Daphnia)

139
Why do predatory invertebrates select for
smaller prey items?
Pastorok (1981)

Chaoborus
has “swim bladders”
sit and wait predator

Daphnia is the prey

Encounter rate between predator and prey
Pastorok (1981)
140

goes up linearly with Daphnia size…

…because
Chaoborus
detect prey by
their wake.

141
But handing time goes up exponentially with size
 142
The relationship between encounter rate and strike
efficiency suggest that medium-size prey are the optimal
diet choice
Encounter rate

Strike efficiency

143
How do these predators find their prey?
1) Wakes
Different sized animals have different sized wakes

Many predators cue into this movement. They are

attracted to small wakes, repelled by larger wakes
 144
How do these predators find their prey?

2) Chemical cues

Many copepods are covered with

chemosensory pits, can “smell” their prey

145
It is not just size that matters, it is overall visibility
Zaret 1972
Two morphs of Ceriodaphnia
Big eye Small eye

Fish always took the big-eye form.

Artificially made small-eye morph more visible by feeding

them india ink. Predation rate increased
Carrying eggs also makes females more visible and 146
increases predation rate Hairston et al. 1983 (L & O)

Sunfish (Lepomis—both bluegill and red breast) feeding on

copepods

Reaction distance
Male Diaptomus 9.2 cm
Female Diaptomus (no eggs) 10.1 cm
Female Diaptomus (with eggs) 12.9 cm

When convert reaction distance to total area searched

(volume of sphere) eggs make females 2x as vulnerable

147
Daphnia pulex with and without ephippia
Melors 1975 (Ecology)

9
8
7
6
number eaten

5
4
3
2
1
0
With ephippia Without ephippia