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This passage is based on the text and images published as:

NORTON, T.A. 2002. And in the beginning …, the pioneering ecological work at Lough Hyne. In: J.D. Nunn (ed.) Marine biodiversity in Ireland and adjacent waters, p. 1-4 Ulster Museum, Belfast

AND IN THE BEGINNING . . .
THE PIONEERING ECOLOGICAL WORK AT LOUGH HYNE

T. A. Norton
School of Life Sciences, Liverpool University, Port Erin Marine Laboratory,
Isle of Man IM9 6JA


Abstract

Now that field experiments are commonplace, it is difficult to imagine a time when it was inconceivable to experiment outside the laboratory in the uncontrollable chaos of the sea. It is even more surprising that the researchers who virtually invented the manipulation of communities in the sea, did marine biology as a hobby. Jack Kitching was the authority on the physiology of the Protozoa, John Ebling a world famous dermatologist, yet they transformed the way we research the shallow sea. In Lough Hyne, a marine ‘lake’ in southern Ireland, they discovered the perfect living laboratory, and the Lough became one of the best-studied and mostsignificant sites in the history of marine biology.


The biological richness of Lough Hyne, a sheltered marine basin in West Cork had long been known (Minchin, 1991). For example, over half of all the seaweed species found in the British Isles have been recorded in the vicinity of Lough Hyne (Norton, 1991), and similar claims could be made for several groups of invertebrates. It was, however, Professor Louis Renouf of University College, Cork who set up the first laboratory there in 1925 (Figure 1). It was in the days when equipping a field laboratory meant buying gallons of preservative and collecting lots of bowls, dishes and other containers. A cardinal sin was to throw away a jam jar. Renouf set about recording the fauna of the Lough and advertising its potential for research (Renouf, 1931). This attracted many distinguished researchers, but surprisingly the most important proved to be parties of students from Bristol University in 1938 and 1939. After the war they returned to begin the concerted study of the ecology of the Lough.

renoufkitching ebling

Figure 1. Louis Renouf, 1930s. Figure 2. Jack Kitching 1956.© Brian Selman Figure 3. John Ebling, 1950s.

Two of the party became internationally famous in their fields: Jack Kitching was probably Britain's leading protozoologist (Figure 2), and John Ebling was a renowned endocrinologist who edited the textbook of dermatology (Figure 3). But for the next forty years, their hobby remained the ecology of Lough Hyne. Every year they returned to spend the summer at the Lough. Initially they used Renouf's Laboratories, but later Jack Kitching, who had private means, built his own. They brought with them parties of students from Bristol University, and later from the University of East Anglia - not on a conventional field course, but to participate in real research. They were the ideal team: Jack (the ideas man) made it happen; John (the organiser) made it work, and the students provided the hands. Kitching appreciated that the Lough was a wonderful living laboratory in which nature had carried out a collection of natural 'experiments', and all he had to do was to monitor the results.

They began with the Rapids, the narrow channel through which the sea rushed in and out of the Lough. What better demonstration could there be of the effects of water flow on the distribution of organisms?

They initially mapped the Rapids and the distribution of its inhabitants, and measured the velocity of flow at different depths throughout the channel over entire tidal cycles (Bassindale et al. 1948). At first they tried to anchor a rowing boat in the stream, but it was swept away and capsized in the cataract. Jack and John only just made it to shore. From then on they used a dispensable student with a current meter in the boat, while the rest of the party, deployed on either bank, fought a tug-of-war with the current to keep the boat stationary. The indirect effects of flow such as sediment and temperature differences were also studied. The sediment was dried on an oven perched over a primus stove.

Jack flung himself in to the work - and the water - wearing a primitive diving helmet that still looked suspiciously like the milk churn it had once been. While others, on the surface, laboured over the air pump (Figure 4), Jack tirelessly hauled up boulders so that their fauna and flora could be quantified (Lilly et al. 1953). He also devised a miniature diving bell in which boulders could be retrieved with their sediment undisturbed. Water samples were taken from numerous locations and depths, and both salinity and oxygen content were laboriously determined with wet chemistry. The party worked in shifts with each group alternating between three hours of continuous sampling in the field and three hours analysing the samples in the laboratory. It never crossed Jack's mind to schedule a rest period in between. In addition to the laborious determinations of pH, 250 water samples were taken, processed and titrated. Starting at four in the morning they measured, weighed and titrated until dusk. Ebling (1991) has commemorated their gargantuan efforts.

rapids

Figure 4. Ronald Bassindale telephones the submerged Kitching and a student pumps down air.

They then turned their attention to what became of the water that entered the Lough. How far did it penetrate and mix with the Lough water? Currents were tracked with floating oranges and sealed milk bottles filled with foxglove flowers to make them more conspicuous. The currents below were followed by means of huge vaned homemade buoys that could be sunk to any required depth. Baby's feeding bottles were also used. They were filled with warm jelly and then deployed at different depths tethered to a rope. In the cool water the jelly began to set, but the current deflected the bottles and the jelly set at an angle. The angle reflected the amount of deflection and thus the current velocity. A tiny compass floating on the jelly indicated the direction of the flow.

To determine the amount of mixing during inflow of water from the sea, John Ebling upset into the Rapids large tubs of water in which 4kg of fluorescein had been dissolved. He managed this by rocking the boat (something at which he excelled), and did it so convincingly that the boat capsized and he had to swim for it in a green cloud whilst he gradually changed colour. The locals, who had heard that the 'mad doctors' were to attempt something spectacular, lined the cliff above and cheered. The dye swirled into the Lough where boats had been moored at strategic positions. Each had a spaghetti tangle of rubber tubes suspended at different depths. For three hours water was sucked up with old Air Raid Precautions stirrup pumps so that the concentration of dye could be determined with yet more titrations (Bassindale et al. 1957). Thus, thanks to Jack's flair, the path of the water and the degree of mixing were determined with equipment that every other household in Britain was throwing out now it was no longer needed for dousing incendiary bombs.

There were more 'natural experiments' awaiting their attention. A 90-metre-deep cave in which the horizontal zonation of seaweeds mirrored their vertical zonation in the water outside, offered an opportunity to differentiate the effects of the progressive dimming of light from the additional changes in wavelength that occurred with depth (Norton et al. 1971). Similarly, in the deep (over 50 metres) Trough running down the western side of the Lough, a thermocline developed every summer, and below, a progressive depletion of oxygen wiped out almost the entire bottom community. It stimulated a long-term study of the resilience of a community exposed to annual devastation. The Trough was monitored like a patient in intensive care. The water temperature at different depths was taken every hour, and oxygen profiles too. At the same time the fate of the benthic fauna was revealed by means of 96 grab samples taken throughout the summer (Kitching et al. 1976). This was a mammoth task of sieving and sorting - one sample alone contained over 600 worms and 5,000 tiny snails. Jack never stinted on samples. Not surprisingly, several projects, including this one, took over a decade to complete.

Lough Hyne was becoming the most intensively studied patch of sea water in the world. But by now Kitching and Ebling were looking at the ecology of the Lough from an entirely different perspective (Figure 5), and this was to transform marine ecology.

kitching
Figure 5. Kitching emerges from just below the Rapids wielding an enormous sickle, with which he has collected a kelp.

It began with a simple observation. Mytilus, the common mussel was abundant in the most sheltered, almost stagnant, parts of the Lough and on the wave-bashed shores outside, but uncommon everywhere in between these two extremes. To find out why, they did what now seems obvious, but was not so then. They transplanted the mussels into the intermediate locations, and found that they were rapidly consumed by crabs and starfish - predators that were uncommon at the more extreme sites (Kitching et al. 1959). It was a striking example of the distinction between a potential and realised niche, and convinced them that it was not the physical and chemical conditions that were overidingly important in the shallows; often it was the predators that were calling the shots. Thereafter much of the work at Lough Hyne became a search for those key organisms, predators, herbivores and competitors, who were determining the structure of the communities. Methods were devised to goad ecosystems into revealing how they were organised. Selected species were added or subtracted, or their abundance was adjusted, or they were transplanted to areas of greater or lesser risk, or made to confront each other within the confines of a cage. In short, they invented experimental marine ecology.

The other great benefit of closely studying the same site year after year was that we (for by this time I was part of the team) noticed any changes that had taken place. For example, in the early years you could not step in the shallows of the Lough without being spiked by a sea urchin. Paracentrotus lividus was widespread and abundant. But in 1971 there were far fewer of them. We confirmed this by counting all the urchins in the southern basin of the Lough and comparing this with similar counts made only a few years earlier. The numbers had fallen from 35,000 to 3,230 and fell even further in subsequent years. This was attributed to crabs whose number had increased ten-fold in the same period - again we had counts of crab numbers taken in earlier years using standardised, repeatable methods (Kitching & Thain 1983). Urchins transplanted into 'barren' areas, where once they had flourished, were demolished by marauding crabs within a few nights. It seemed that in the past too there had been cyclic changes in the relative abundance of all these organisms.

A consequence of the decline in grazing urchins was the blossoming of the green alga, Codium, which now formed a bright swathe in the shallows. But how could its abundance be quantified? Nowadays we would employ aerial photography, and interpret the pictures with image analysis and, of course, consume a large grant in the process. Jack Kitching's genius was that he could do it just as well using a rowing boat, a long ruler, a net bag and a spring balance. The total cost of all the equipment including the boat was less than £50.

Sadly, the decline in the urchins now appears to be permanent rather than a short-term phenomenon. The temperature in the Lough seems to be changing, - perhaps, according to David Barnes, it is yet another consequence of the El Niño Southern Oscillation. Whatever the cause, the urchins are now uncommon in the Lough and have been so for several years.

I first came to the Lough as a young research student in 1964. I arrived by water, having no alternative but to swim to the isolated laboratory site. I rose from the waves festooned with seaweed to be confronted with a tall figure in a sweater that had clearly been savaged by sharks.

"Professor Kitching, I presume?"
"King Neptune without a doubt," he replied.
"Welcome to the Glannafeen Laboratory."

It was to be the first of fourteen wonderful summers that I was privileged to spend in the company of Jack and John, learning to be an ecologist. I also discovered what could be achieved with the minimum of fancy equipment and the maximum of ingenuity.

Thanks to Kitching, Lough Hyne was declared the first statutory marine reserve in Europe. Although both Jack and John are now dead, the work continues. They would be delighted that Lough Hyne is still the focus for researchers from the Universities of Bangor and Newcastle and especially Cork. Jack donated his laboratories to U. C. Cork, and they have since built a new one of their own. A dozen projects are underway, and it is no longer predominantly a summer camp; research goes on all year round. Best of all, new generations of students continue to come to the Lough and will inevitably fall under its spell, just as they have been doing for almost 80 years.

Those who wish to read more of the ecological work of Kitching and Ebling at Lough Hyne should begin with their review papers (Kitching & Ebling 1967; Kitching 1987). My own book, Reflections on a Summer Sea (Norton 2001), attempts to capture the experience of working with two such extraordinary people and doing research just for the fun of it, in the days when it was all mad, magical and marine biological.

Copyright © Trevor Norton