Sei sulla pagina 1di 15

Population Growth and Competition in Lemna sp. and Spirodela sp.

Daniel Seth Andal

BIO 150 F-1L

A Scientific paper submitted in partial fulfilment of the requirements in BIO 150 (Ecology)
laboratory under Prof. Kristine O. Abenis, 1 st semester, 2019-2020.

Population growth and competition were observed among Lemna sp. and Spirodela sp. The
fronds of both species were placed in pure and mixed cultures with two treatments for each
culture as it is observed ten times every other day. The results show that Lemna has greater
population when subjected to the nutrient treatment and Spirodela on distilled water
treatment. The results from the mixed culture shows that both can coexist with each other
as the population is greater at the treatment with soil in its environment. The data shows an
exponential increase in population growth and some minimal decrease at some point which
regards that population size can exceed carrying capacity but will stabilize itself below the
maximum requirements if exceeded. Graphs constructed from these data shows a logistic
growth curve which can be supported by the computed instantaneous growth rate between
intervals of observation days. It is concluded that Lemna is the more competitive and fit
species of duckweed due to its proportional growth with the concentration of nutrients in
its environment, greater population size on distilled water medium, and morphological
advantage over the other.

Population growth is defined as the increase and decrease in the number of

individual with time in a certain population. In this experiment, the population growth of

Lemna sp. and Spirodela sp. were focused on as it is treated into their respective ecological

communities. Lang and Benbow (2013) defined ecological community as an assemblage

of populations of at least two different species that interact directly and indirectly within a

defined geographic area. The two species of duckweeds are floating macrophytes which

competes for space, light, and nutrients along a gradient of biotic and abiotic factors.

Harper (1977) suggests that these types of free-floating aquatic plants are excellent models

for growth rate determination due to its continuous growth compared to most plants that

have life cycles with discrete jumps in their population size such as alteration in size and

shape as response to change in population density.

Kopeny (2002) explains that a population growth rate has three phases when

resources are depleted. It is known as logistic population growth where growth rate slows

and eventually stops, thus, exhibiting a sigmoidal or S-shaped growth curve which can be

observed at Figure 1 below.

Figure I. A Logistic Growth Curve

Interspecific competition occurs between two different species when the resources

are limited at their disposal. The Gause Competitive Exclusion Principle states that two

species with similar ecological requirements cannot coexist and will undergo competition

with each other. Volterra (1931) emphasizes that 4 outcomes of these interactions can be

predicted by the Lotka-Volterra model whereas it can be one of the species prevail and the

another would eventually be completely excluded, and vice versa, either species wins based

on population density (unstable equilibrium), or coexistence will occur. In addition to the

first two outcomes, Jefferies (2000) stated that two species sharing limited resources may

result to depression of population growth rate and maximum population size of either of

the species. To distinguish Lemna sp. from Spirodela sp, it is relatively smaller but both

species have a circular thalli or fronds. The hypothesis: Lemna sp. will have higher

population on both treatments in pure and mixed cultures than Spirodela sp. can be


The main objective of this study is to determine the changes in the population

growth and competition of Lemna sp. and Spirodela sp. The specific objectives were the


1. to determine the effects of intraspecific and interspecific competition on

both species;

2. to compare the growth of Lemna sp. and Spirodela sp. in different types of

media; and

3. to demonstrate changes in the populations of pure and mixed cultures of

Lemna sp. and Spirodela sp.

This exercise was conducted on 19th of August 2019 to 11th of September 2019 at

the Institute of Biological Sciences in the University of the Philippines Los Baños.


Three replicate jars were prepared for treatments A and B whereas treatment A

contains 50 mL of distilled water and treatment B contains 3 g of soil and 50 mL of distilled

water. Ten fronds of Lemna sp. were introduced to one set of treatments A and B. The same

number of Spirodela sp. fronds was also introduced to another set of similar treatments.

Separating the fronds during the preparation of the pure cultures was avoided to distinguish

the count easily and represent the initial population (N o) as 10. Five fronds of Lemna sp.

and another five fronds of Spirodela sp. were introduced to the remaining sets as these

represent the mixed culture where the No is also equal to 10.

After preparing the cultures, the jars were covered with transparent plastic and was

punched few small holes to give aid for aeration. These cultures were kept in the designated

area in a greenhouse and assured that all fronds are floating in the media.

The total number of fronds of Lemna sp. and Spirodela sp. were counted every

other day for each treatment until the frond count levels off. Visible new buds that are half

the size of the old buds were counted as separate individuals while the fronds that turned

yellow or dried up were disregarded in the count. After each observation, it is assured that

no fronds are left adhering on the sides of the bottle so that all fronds are still floating on

the media. This is done by minimizing the shaking of the bottles.

The data obtained after 10 observations were plotted in graphs where it is separated

for each treatment. Frond count VS time was plotted as separate line graphs for Lemna

grown as pure and mixed cultures. Another two graphs follow the same attributes for

Spirodela. The instantaneous growth rate (rinst) at each observation interval in all cultures

were computed using the following formula:

𝜆= (Equation 1)

rinst = ln λ (Equation 2)

where Nt = population time in t

N(t+1) = population at the next observation day
λ = annual finite rate of increase
rinst = instantaneous growth rate

After 10 observations, the results from the obtained data is as follows for both

media for both cultures:

Table 1. Population Growth of Pure and Mixed Culture of Lemna sp. and Spirodela sp.

Age of Lemna sp. Spirodela sp. Lemna sp. + Spirodela sp.

Date Culture A B
# A B A B
(days) L S L S
1 8/19/19 0 10 10 10 10 5 5 5 5
2 8/22/19 3 10 10 10 10 5 5.67 5.33 6
3 8/27/19 8 16.67 16.33 20 12.33 6.67 8.33 5 7.67
4 8/29/19 10 19.33 19.67 23.33 16.67 11 11.33 12 11.33
5 8/31/19 12 20 20.33 24.33 16.33 13 11.67 14.67 11.67
6 9/2/19 14 22 22.67 27.33 14 13.67 13 17 13.67
7 9/4/19 16 22.67 23.67 29 15.67 15.67 14.33 22.33 13
8 9/6/19 18 27.33 31.67 28.67 22.33 17.33 15.67 30 15
9 9/9/19 21 30.33 34 30.67 25.33 21.33 16 32.33 20.33
10 9/11/19 23 32.33 36 32.33 26.67 23.33 18.67 33 21
Legend: A – dH2O medium; B – dH2O + soil medium

Table 1 shows the mean population of the Lemna and Spirodela throughout the

course of 10 observation for 23 days. This shows the trend of the duckweed growth over

the mean of three replicates for each treatment. Generally, Lemna has greater population

density than of Spirodela. It can be observed that in pure culture, Lemna has greater

population size when subjected to distilled water and soil medium but on the other hand,

Spirodela’s population size is less when subjected to a medium with nutrients. This effect

on Spirodela’s population size is an example of intraspecific competition wherein

competition occurs with members of the same species in the population. According to the

Centre for Ecology and Hydrology (2004), Lemna grows best in waters mixed with

different nutrients. This explains the trend of Lemna having greater population density on

the soil and distilled water medium where its environment is eutrophic. On the other hand,

the effect of nutrients to Spirodela tends to decline its growth. Liu (2017) explains that
Spirodela growth increases only at media where nutrient concentration is at its smallest.

Additionally, he emphasized that metal compounds concentration increases along with

nutrient concentration which hinders the reproduction and growth of Spirodela.

For the mixed culture, it can be observed that Lemna is still greater in population

density than Spirodela on either type of medium. Although it is discussed earlier that

Spirodela has lower population density when subjected to the medium with nutrients, the

results for the mixed culture shows the opposite. This is a possible result of these two

species undergoing coexistence. A related study by Bauer et al. (2008) compared

population growth in competing species of Salvinia and Spirodela, the research concludes

that factors coupled with mosaic of wetland environments promoted a continuum of

competitive outcomes between the two species and eventually ensured their continued

coexistence. This environment is similar to the pure culture set-up B and the coexistence

of Lemna and Spirodela is evident as their population growth thrived higher than of the

distilled water medium.

Table 2. Instantaneous Growth Rate of Lemna and Spirodela between day intervals

The instantaneous growth rate between day intervals of the observations shown in

Table 2 were computed to monitor the increase and decrease of the population between
observation days. It is evident that the last observations at 9-10 days shows stabilizing

values and significantly lower than those of between the previous observations. This agrees

with the logistic growth curve which is illustrated at Figure 1. In pure cultures, it is

observed that Spirodela decreased in population 6 times, since there are 6 negative values

obtained from the calculated instantaneous growth rate, and Lemna only exhibited 3 times

of population decline. In mixed cultures, Spirodela also had more population decline, with

5 negative values, significantly on the nutrient medium, and Lemna only had 3 negative

values for the computed growth rate. This shows that the biomass of the species has already

reached its carrying capacity. However, it is restored again with positive values for the next

interval which would mean that the logistic growth curve would be followed with medium

time lags and dampened oscillations. Additionally, Cuevas et al (2012) suggests that the

growth of the population will be negative, indicating a population decrease until the

population size and the carrying capacity will become equal and stable.

Figure III. Lemna in Distilled Water Medium Figure II. Spirodela in Distilled Water Medium

Figures 2 and 3 shows the population growth curve of Lemna and Spirodela,

respectively, in distilled water medium. It resembles a sigmoidal or S-shaped curve which

conforms to the logistic growth curve illustrated in Figure 1 and the computed data for the

instantaneous growth rate. It is thorough for both species that pure culture provided greater
population growth than mixed culture. This occurrence in mixed culture was a result of

differential resource utilization. Since it is both in distilled water medium, the only

resources these species could utilize among the mixed culture is sunlight and space. The

population growth is lower on the mixed culture but coexistence occurs since the combined

resource consumption of the competing species equals its resource supply.

Figure V. Lemna in Soil + Distilled Water Medium Figure IV. Spirodela in Soil + Distilled Water Medium

Figures 4 and 5 above shows the difference of pure and mixed culture when the two

species of duckweed is subjected to soil in distilled water. The results can be observed to

also have a sigmoidal or S-shaped curved which also indicates that it is a logistic growth.

Although the results resembles the one with distilled water medium with pure culture

possessing the greater frond count, it is evident that the curves touched or obtained

relatively close values at certain points. This is a result of resource partitioning through

differential resource utilization in which one species uses a portion that is less used by the


Carrying capacity is defined as the number of individuals from a population that

the environment can support. The pure culture with soil and distilled water media provided

the higher carrying capacity for Lemna. This is the result of its growth rate being

proportional to the nutrient availability as Farmer (2016) has explained for this genera. On
the other hand, the pure culture with distilled water media provided the higher carrying

capacity for Spirodela as earlier discussed, that higher nutrient concentration depletes the

growth of Spirodela since it is reinforced with higher metal concentration which affects the

growth of Spirodela negatively, thus, it has higher carrying capacity on a medium without

soil and its minerals and nutrients within.

A well-established sigmoidal curve that represents logistic growth among the other

set-ups is the one with Spirodela on distilled water medium. The exponential phase and

stationary phase can be clearly distinguished in the graph on Figure 3. The instantaneous

growth rate values reinforce this statement as it has relatively higher and outlier values on

the observations with increasing trend and has relatively close values during the final stages

of observation. To apply the phases of population growth, 0-3rd day is the lag phase, 3rd-

12th day is the exponential phase with an emphasis on the spike between the 3 rd- 8th day

interval and the 12th-23rd day represents the diminishing growth phase with only a few

fronds being observed to be added to the population.

The growth curve that is observed between the pure and mix cultures appears to be

distinct with each other as mix culture exhibits a lower population growth than of the pure

culture. This is affected by changing birth and death rates which can also be observed in

Table 2 where there are decreasing population between intervals. Additionally, there are

density-dependent factors that affect these curves such as competition between the two


Lastly, the effect of competition to these species is that their population growth is

lower. It can be observed that pure culture has higher population size than mixed culture

but it is evident that in the mixed culture with soil and distilled water medium, the
population is higher for both species than the medium without nutrients. As mentioned

earlier, this is an effect of resource partitioning through differential resource utilization.

Bell, Stambolie, and Leng (1995) explains that nutrients are absorbed through all surfaces

of the duckweed leaf. Since Spirodela has larger leaf size than Lemna, it has a tendency to

absorb nutrients more than the latter, but it is mentioned that higher nutrient concentration

decreases its population growth. Hence, Lemna will have the better competitive ability.

Duckweed population growth is not only controlled by the nutrients it receives but also by

its space and sunlight resources. Since Lemna is smaller, it would occupy less space and

will have more for its reproduction. Additionally, Keddy (1976) argues that Lemna is more

competitive and fit than other species under the family Lemnaceae since it has the ability

to increase its carrying capacity along with increasing nutrients and high tolerance to

different pH ranges.


Duckweed species of Lemna sp. and Spirodela sp. were subjected to be observed

in terms of their population growth and competition. The set-up involves two cultures, pure

and mixed, with two treatments for each. Treatment A contains distilled water and

treatment B contains soil and distilled water. These were put in jars and observed every

other day until 10 observations were made. In the pure culture, Lemna has a higher

population on treatment B while Spirodela has higher population on treatment A. This is

reinforced by the explanation that Lemna has proportional growth rate with the

concentration of nutrients in its environment while Spirodela grows best at environments

with the smallest amount of nutrient concentration. In mixed culture, Lemna is evidently
greater in population size in both treatments, but it is significant that the individual

population of both species is greater on the medium with soil and can be explained through

resource partitioning through differential resource utilization. The calculated instantaneous

growth rate conforms to a logistic growth curve being sigmoidal or S-shaped.

Taking everything into account, the hypothesis: Lemna sp. will have higher

population on both treatments in pure and mixed cultures than Spirodela sp. is accepted.

Spirodela having larger fronds consumed much of its space and relied mostly on the

sunlight as a resource and Lemna having proportional growth to the nutrient concentration

of its environment makes it a more competitive species than Spirodela. It is certain that the

carrying capacity of a population can be exceeded and will undergo negative instantaneous

growth rate after the said overrun. Taking all these data into application, it is still needed

to engage in different speculations to be validated by further studies. Other species of

duckweed can be taken into account, adjusting of its environmental factors such as pH and

mineral content. The possible errors in this experiment is the availability of sunlight as a

resource in the storage place for the set-ups. It is also possible that the jars were shaken

which may have caused the death of some of the fronds. It is recommended to use the same

amount of time since the logistic growth curve can be easily distinguished over the data

gathered from 10 observations.


BAUER, L., CENTER, T.D., MARTIN, M.R., TIPPING, P.W. 2008. Competition
between Salvinia minima and Spirodela polyrhiza mediated by nutrient
levels and herbivory. Aquatic Botany. Journal retrieved from pp. 231-234.

BELL, R., LENG, R.A., STAMBOLIE, J.H. 1995. Duckweed - a potential high-protein
feed resource for domestic animals and fish. Livestock Research for Rural
Development. Centre of Duckweed Research & Development University of
New England Armidale, NSW 2351. Volume 7, Number 1.

CENTRE FOR ECOLOGY & HYDROLOGY. 2004. Information sheet: Lemna species.
Retrieved from

DUPO, A.L.A. & LAMBIO, I.A.F. 2012. Exploring Ecology (no ed.).
University of the Philippines Los Baños: Environmental Biology Division,

FARMER, J. 2016. Effects of Nutrient Availability on Growth Rate Capacity of Lemna

minor. Boise State University.

HARPER, J. L. 1977. Population Biology of Plants. Academy Press, NY.

JEFFERIS, R.L. 2000. Population Growth: Experimental Models using Duckweed (Lemna
sp.). ESA lab pp. 1-3.

KEDDY, P. 1976. Lakes as Islands: The Distributional Ecology of Two Aquatic Plants,
Lemna Minor L. and L. Trisulca L. Ecology 57:353-359.

KOPENY, M. 2002. Lecture #K5 - Population Ecology, Continued. In BIO 202. Retrieved
LANG, J. M. & BENBOW, M. E. 2013. Species Interactions and Competition. Nature
Education Knowledge 4(4):8.

LIU, C., CHEN, L., LIANG, X., JIN, L., SHI, H., XIANJIANG, K., and, ZHAO, Z. 2017.
Inter- and intra-specific competition of duckweed under multiple heavy
metal contaminated water. Aquatic Toxicology. DOI:

VOLTERRA, V. 1931. Variations and Fluctuations of the Numbers of Individuals in

Animal Species Living Together. In: Chapman, R.N. Animal Ecology.
McGraw-Hill, New York.