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Population dynamics: Phytoplankton

1. How do phytoplankton populations change over time?

2. How do we calculate population growth rate?

3. What are the major biotic and abiotic factors influencing phytoplankton dynamics?

Significance Algae are food for zooplankton, so algal dynamics influence zooplankton dynamics. Zooplankton are food for fish... Overabundant algae can be a nuisance

Basic ecological question: what controls distribution and abundance of species?

Some terms to keep straight:

primary productivity: is a rate that describes the production of organic carbon (e.g., mg C/m2/day) r: population growth rate. Describes the rate of change of N (e.g., numbers/day) standing stock: the amount of algae (or other organic material) present in a lake at any time. Reflects both the productivity and the turnover. euphotic zone water layer with sufficient light for photosynthesis ( epilimnion)

How are these things quantified?

Productivity is often measured on isolated samples as a change in oxygen concentration or as uptake of radioactive carbon (14C) into algae
Population growth rate (r) is measured over time by counting cells of a particular species Standing stock can be measured with cell counts or by chlorophyll a extraction All these things change over time

Typical pattern in temperate, dimictic lake

Phytoplankton Biomass (g/l)

6000 5000 4000 3000 2000 1000 0

spring bloom fall bloom


ar A pr M ay Ju n

See temporal changes in both abundance and species composition (seasonal succession)

Ju l A ug

Se p O ct N ov D ec

n Ja


Algae reproduce by cell division

N2 = 4
N0 = 1 populations double every generation often about 12 48 hours for algae N1 = 2

N3 = 8

Can measure the rate of population growth

Assuming generation time = 12 h, and no death

300 250

Time (hours) 0

Cell count 1

r (per hour) 0.06

cell count

200 150 100 50 0 0 50 100 150

24 36 48

4 8 16

0.06 0.06 0.06

72 84 96

64 128 256

0.06 0.06

time (hours)

But, no population increases forever

How fast is the population size changing (r)? Can measure in the field or lab


ln Nt ln N0 t

change in N change in time

N0 = population at time 0 Nt = population at time t t = time If: r > 0, population is increasing (growing) r = 0, population size is not changing r < 0, population id decreasing (shrinking)

Why do populations increase? Birth, immigration

Why do populations decrease? Death, emigration If we ignore immigration and emigration: r=b-d

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

If: r > 0, there are more births than deaths r = 0, births equal deaths r < 0, there are fewer births than deaths

Why do births and deaths change over time?

Loss #1

Grazing rateloss due to feeding by zooplankton Variation in how easily things are eaten by zooplankton. Loss rate depends both on the algae and on the grazer
http://dnr.metrokc.gov/wlr/waterres/lakes/daphnia.jpg www.uv.es/~ciros/zoopl_en.html http://www.eeb.cornell.edu/hairston/hairston.html

Brachionus (300 m)

Diaptomus (< 2 mm)


(2-3 mm)

Cladocera, especially Daphnia, are the main grazers in aquatic systems.

Loss #1 Tessier et al. (2001) and many other studies have shown:
Lots of Grazing on:
Single celled green algae

Small Diatoms

microbes.limnology.wisc.edu/outreach/ images/peridinium.gif

Chlamydomonas Not as much grazing on:




Gelatinous greens




Loss #2

Sedimentation ratesinking out of the euphotic zone How will time of year and mixing pattern of a lake influence algal loss by sedimentation rate?

Sedimentation rate is estimated by measuring the rate of sinking Vs and comparing that to the average depth of the epilimnion Sinking rate can be from 0.1-6 m per day in a calm water column

Loss #2 Sedimentation ratesinking out of the euphotic zone

Algae avoid sinking by:

1. Increasing surface-to-volume ratio


Loss #2 Sedimentation ratesinking out of the euphotic zone

Algae avoid sinking by:

2. Mucilaginous or gelatinous sheath (reduces density)


Loss #2 Sedimentation ratesinking out of the euphotic zone

Algae avoid sinking by:

3. Forming gas vacuoles (common in blue-greens)
www.biologie.uni-hamburg.de/ b-online/fo42/anabaena.jpg


Loss #3 Physiological mortality (things die) and mortality due to parasites Traditionally, this loss rate was assumed to be small

But, that thought is slowly changing as more research focuses on disease.

Rhizophydium granulosporum (parasite) on Oedogonium (host)

Loss #4

Washout loss due to outflow Can be estimated from ratio of outflow to volume of epilimnion

Usually unimportant unless there is flooding

What influences the rate at which cells divide (b)? (1)Temperature: In general, cells divide faster in warm water, but relatively speaking... Diatoms grow better in cold water

Blue greens in warm water (2) Competitive ability

Cells cant divide until they have enough energy, and different species are better or worse at acquiring limiting nutrients

Also... Mixing is important for diatoms, because they are heavy.

Diatoms are most abundant under conditions of circulation (spring and fall in a dimictic lake).

Mixing pattern also influences nutrient cycling, abundance of potentially limiting nutrients

Summary of phytoplankton dynamics: Gains (b) depend on: Temperature & Mixing Losses (d) depend on: Temperature & Mixing Washout loss due to outflow Grazing Resources (competition)

Physiological mortality and mortality due to parasites

Concepts to know
Understand the difference between primary productivity, r and standing stock. Know the typical seasonal pattern for phytoplankton abundance. Know how to calculate r and what this number tells you. Understand the primary factors contributing to algal death rate. Are these factors of equal importance for all species?

Practice question
You have been sampling populations of Cyclotella comensis, and Oscillatoria limnetica from Lake Michigan. Data are presented below. Date 1-Apr-04 8-Apr-04 1-Aug-04 8-Aug-04 # of Cyclotella cells/ml 10 2600 35 15 # of Oscillatoria cells/ml 170 180 1200 2500

Which species has the highest growth rate in April? In August? What biotic and abiotic factors likely contribute to the temporal differences in growth rate and abundance?

Zooplankton population dynamics

1. What are the similarities and differences between the growth of algal populations and zooplankton populations? 2. How can we use the Edmondson egg ratio method to understand the dynamics of natural populations of zooplankton? 3. What is DVM?

For rotifers and cladocera, most of the time reproduction is by parthenogenesis The seasonal dynamics can look like the figure for phytoplankton:

Remember: r=b-d

Why did the population decline? 1. b decreased and d remained constant 2. b remained constant and d increased 3. b decreased and d increased

In natural populations, how can you tell?

In zooplankton, gains (birth rate-b) are influenced by: Temperature Resources Losses (death rate-d) are influenced by: Predation Physiological mortality and parasites Washout losses due to outflow Unlike the phytoplankton, sedimentation is not a loss term for zooplankton because they can swim

When you sample a population regularly, you can estimate the number of animal/L on each date.

From this, you can calculate the population growth rate:


ln Nt ln N0 t

From the samples, can also count the number of eggs carried by each female (clutch size)
Brachionus Daphnia ephemeralis


In rotifers, clutch size is usually <3

In some big


maximum clutch size can be ~ 100

In well fed cladocera, clutch size is correlated with body size.

Once you know the population growth rate, and the number of eggs/female (egg ratioE), can begin to address factors regulating population dynamics. Edmondson egg ratio method (1968)
b= ln (E + 1) D

E = number of eggs / female D = development time of the eggs (hours or days) For both rotifers and cladocera, development time is temperature dependant. Eggs develop faster at warmer temperatures

Development time for various species of cladocera

Assumptions of the Egg Ratio Method: Stable age distribution of the eggs Closed population


ln (E + 1) D

ln Nt ln N0 t

Can solve for d

Dudycha 2004

Can then use seasonal variation in birth and death rates to address changes in strength of competition vs. predation (or parasites)

It is possible to get negative death rates, what does this tell you?

d (day


b (day -1)

r (day -1)

For rotifers and cladocera, the relative importance of food limitation (competition) vs. predation (or parasites) can be determined from preserved samples. The egg ratio method works when: 1. Females are reproducing rapidly 2. Animals have short generation times 3. The investigator samples frequently The egg ratio method also assumes that reproduction is asexual, it ignores the annual period of sexual reproduction

Copepods are different:

Always reproduce sexuallyneed to find mates

Have much longer generations times

Instead of constant reproduction, have cohort structure

Cohortgroup of individuals all born at the same time

Since copepods have morphologically distinct instars, can conduct instar analysis

Generation times for copepods are much longer than for rotifers or cladocera

In many lakes, copepods have one or two generations per year, whereas rotifers can have 40 (cladocera are in between)

How might this difference in generation time influence interactions among species?

Population dynamics are influenced by the behavior of individuals Individual behavior: How to gain enough food and mates to produce the most offspring while avoiding death as long as possible?

Minimize costs while maximizing benefits

The environment is constantly changing

There are structural and physiological limits to adaptation

Predator avoidance behavior:

Diel Vertical Migration (DVM) Daily change in the distribution of individuals in the water column

Typical pattern is to find large zooplankton in the hypolimnion during the day and observe movement into the epilimnion at night

Distance of migration can be a few meters to over 100 m

Influenced by:

Developmental stage or gender

Transparency of the water Temperature Food quality and quantity

How do you know where the zooplankton are in the water?

www.aquaticresearch.com/plankton_trap/ Mvc-005f.jpg

Closing Net

Schindler Trap

Why do zooplankton migrate? Proximate vs. Ultimate factors Proximate factors are light and chemical substances (kairomones)

But changes in light intensity do not directly influence b or d

Ultimate factors include predation, food, and egg development time

High predation by fish High UV exposure Higher temperature Low predation by inverts. High food (sometimes) Low predation by fish Lower food (sometimes) Low temperatures Higher predation by inverts.

Initially, it was believed that food quantity and quality was so much higher in the epilimnion, the grazers must be entering the epilimnion at night to feed.
Recent experiments by Kessler (2004) and Lampert et al. (2003) have suggested a different explanation...

New thought is that sometimes food is better in the hypolimnion, but the grazers enter the epilimnion for the temperature benefit (egg development).

How to test this new idea?? The plankton towers of Pln

11.2 m high 0.86 m inner diameter

The plankton towers have also been used to test the role of fish kairomones in DVM

Vertical profiles of Daphnia hyalina

In lakes where the bottom becomes anoxic, large zooplankton have less of a refuge

In lakes where the entire hypolimnion becomes anoxic, large zooplankton have no refuge and are often eliminated by late summer.

So, population dynamics in zooplankton are influenced by: Temperature things happen faster when it is warmer Species parthenogenesis or sexual reproduction? Individual behavior to migrate or not to migrate? Basin shape and physical limnology - modifies everything else Presence or absence of predators, parasites, competitors

How does the best strategy become dominant in a population? Populations are made up of different genotypes, which differ in their behavior (and other life history traits). When there is variation among genotypes in some trait, and this variation is heritable, there can be differential survival and/or reproduction. If conditions change dramatically, new best strategy can dominate

Concepts to Know

What factors influence r in zooplankton populations? What determines clutch size in zooplankton? What use is the Edmondson egg ratio method to limnologist? What is a trade-off? What types of trade-offs do zooplankton face? Why migrate?

West Gull Pond is a small temporary pond in southern Michigan. The pond typically fills in early March and is dry by July. The pond contains many typical temporary pond species, including a population of Daphnia pulex. During 2004, the pond was sampled at regular intervals from the day it filled on March 15 until it dried in July. At each sampling time, water temperature was recorded, and samples of Daphnia were taken to determine density (#/L), birth rate and death rate. The data birth and death rates are provided below.
Sample Date 30 March 15 April 30 April 15 May 30 May 15 June 30 June 15 July Birth rate (b) (per day) 0 0.6 0.5 0.4 0.6 0.2 0 Pond dry Death rate (d) (per day) -0.4 0.1 0.1 0.1 0.9 0.9 0 Pond dry

Practice Question

At which point during the year did the Daphnia density (#/L) begin to decline? How did you determine this?
From the data provided, what is the most likely mechanism leading to the initial decline in density ? How did you determine this?

What is the most likely mechanism causing the negative death rates observed on 30 March?