An experiment carried out by G. C. Gause in 1934 using species of Paramecium, a large protozoan
common in fresh water. It feeds on plankton, the food source used in these experiments.
food vacuoles of
waste disposed of bacteria formed here
gullet (’cytopharynx’)
products of digestion
absorbed into cytoplasm
P. aurelia
a smaller,
species cultured separately fast-growing species
200
numbers of Paramecium per
0.5 cm3 of culture solution
150
100
P. caudatum
50 a relatively large,
slow-growing species
0
0 2 4 6 8 10 12 14 16
time in days
200
numbers of Paramecium per
0.5 cm3 of culture solution
150
0
0 2 4 6 8 10 12 14 16
time in days
We can see from the graphs that ‘competitive exclusion’ may be the pressure that causes related
species living in close proximity to evolve separate niches. In this way, competition between
them is at an end. Of course, this would only occur over an extended period of time. So, the
competitive exclusion principle would, in the long term, account for the difference in niches
between the two species of paramecium, for example.
Since it was first proposed, the competitive exclusion principle has been demonstrated
convincingly in laboratory studies, but the demonstration of its operation in the environment is
more difficult.
However, the behaviour of two species of barnacle that live on the seashore appears to be a
case in point. Barnacles are sedentary crustaceans colonising any surface in the inter-tidal zone
on which they happen to settle. But before that occurs, barnacles are dispersed to fresh habitats
by their free-swimming larval stage, which eventually settles on some rocky surface it encounters.
Now one of these two species of barnacle, Chthamelus, is able to withstand prolonged exposure
when the tides recede, but these same conditions kill off the other species, Semibalanus. The
result is a characteristic distribution pattern of barnacle populations; Chthamelus barnacles are
found in the upper inter-tidal zones, but are crowded out from lower zones, and for Semibalanus
barnacles, the reverse is the case. We see here that it is a difference in resistance to periodic
desiccation that has determined where each species will survive best. The two species are
adapted to different environmental conditions.
The degree of exposure determines the distinctive distribution pattern of these two species of barnacle:
Chthamalus Semibalanus
6 plates
Interpretations of
Distribution Tide levels
observations and experiments
adults larvae
C mean high
B spring tide
Chthamalus survives
Semibalanus dies through desiccation mean high
neap tide
Semibalanus outcompetes
Chthamalus for space
mean tide
The earliest, significant contribution to acceptance of the ideas of organic evolution was a
geological discovery. It was the realisation that the Earth is extremely old. In Western culture,
the biblical account of creation was generally accepted as authoritative until the eighteenth
century, at least. Furthermore, the chronology detailed in the Bible suggested that life had
appeared on Earth a mere few thousand years ago. This timescale, that the Earth was only
5000–6000 years old, was widely accepted in Europe until well into the nineteenth century.
James Hutton (1726–97), a doctor, farmer and experimental scientist, realised that the
sedimentary rocks of many existing mountain ranges had once been the beds of lakes and seas
and, before that, had been the rocks of even older mountains. He made no estimate of the age of
the Earth, but he realised that, in contrast to biblical estimates, the Earth’s timescale virtually
has ‘no beginning and no end’.
Now, geologists estimate the age of the Earth as being 4500 million years old, and life as
having originated 3500 million years ago. On these timescales, it becomes possible to imagine
organic evolution by gradual changes.
Arctic Circle
60°
40°
Tropic
of
Cancer
20°
20° Tropic
of
Capricorn
Key
40° tropical rainforest Mediterranean scrub forest
temperate rainforest grassland and savannah
temperate deciduous desert
forest
ice and tundra (treeless
60°
boreal coniferous forest plains of marsh and stones)
(taiga) Antarctic Circle