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Breathing out of the water, a true low-tide stress for limpets?

Limpets can resist desiccation and large temperature variations resulting from their air exposure at low tide. Studying their respiration in air and in water and the calcification of their shell helps understanding their adaptation and their distribution in the intertidal zone.

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Emersion is a considerable physiological challenge for marine organisms living between low and high tide marks. Many of them are unable to move or move very slowly and therefore cannot escape the extreme conditions of low tide, such as desiccation and temperature variations. This is the case of limpets: in Brittany, the main species (Patella vulgata) can withstand temperatures over 40°C and survive for long periods out of the water. It has been shown that they owe this capacity to the water remaining between the body and the shell, which prevents desiccation and brings dissolved oxygen for respiration. But much remains to be learnt about the influence of the alternation of emersion and immersion on their metabolism and on their vertical distribution within the intertidal zone.


The intertidal zone, and particularly its highest part, is an extreme habitat where survival implies deep adaptations from animal and vegetal species



The works presented in this paper addressed these questions in terms of carbon metabolism. This element is provided through feeding and can be used to synthesize organic molecules in the soft tissues of the animal or to provide the energy necessary for life; CO2 is then rejected in the air or the water through respiration. The fixation of calcium carbonate CaCO3 in the shell (calcification) also entails carbon rejection under the form of CO2. Researchers studied calcification and respiration according to the season and the location of the animals relative to the tide. Three- to four-centimeter limpets were sampled from January 2010 to September 2011 6 m, (high-shore) 4 m (middle-shore) and 2 m (low-shore) above chart datum. In the laboratory, the animals were placed in tanks where natural environmental conditions were reproduced. Various parameters related to metabolism were measured along incubation experiments lasting from a few minutes (emerged animals) to 1h30 (immerged animals).

P. vulgata is the main species in Brittany; the smallest individuals (about 1 cm long on this picture) are mostly found in the lowest levels of the intertidal zone.

The three components of the carbone mission budget were estimated: calcification, respiration when immerged and when emerged.

CaCO3 deposition in the shell occurs all year round, but higher rates are observed in summer, when temperature and feeding reach maximum values. When high-shore limpets are immerged, their calcification rate is two or three times lower than that of limpets of lower levels.

Irrespective of their level, the carbon flux emitted by immerged limpets is higher in summer than in winter; water temperature is indeed a good indicator of respiration when immerged. The average hourly emission (instantaneous flux) can be converted into a daily flux when the average duration of immersion at each level is taken into account.

The lower the limpets (thus the longer their immersion), the higher their carbon emitted in water through respiration during one day; at 13°C, the emissions at the middle and low levels relative to the high levels are in a ratio of 3 and 6.5.

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Seasonal variation of instantaneous respiration rates of immerged limpets according to their level


When emerged, limpets still breathe but their CO2 flux is influenced but relative humidity of air, in addition to its temperature. In most combinations of these two factors (from 7 to 32 °C and from 26 to 90%), flux at high and middle levels remains stable during a long emersion (6 hours), whereas at the lowest level it is high during the first hour before rapidly decreasing. The duration of emersion influences daily fluxes: at spring tides, high-shore limpets emit in the air about twice as much CO2 as low-shore ones who have a shorter emersion time.

For all levels and temperatures, the instantaneous rate of respiration is higher in the air than in water.

P. vulgata reaches a diameter of 6 cm; the highest individuals spend most of their time emerged.


At a yearly scale, the total amount of carbon emitted by limpet populations is similar at all levels, but its distribution is very different. High-shore limpets spend underwater a time equivalent to only 27 days/year, and 94 % of their carbon is emitted by aerial respiration; middle-shore ones spend 170 days emerged and emit in the water 24 % of their carbon (65 % in the air); low-shore ones are immerged 314 days/year and reject into the air only 18 % of their carbon. The share of calcification within the carbon budget is very low at high levels (2 %) and increases to 11 % and 18 % at middle and low levels.

High-shore limpets thus have a lesser calcification ability; this suggests that the preferential distribution of smaller individuals in the lower shore may not be related only to their sensitivity to desiccation, but also to their requirements in calcium carbonate for shell growth. Conversely, the larger individuals whose needs are reduced could withstand longer emersions; their vertical migration pattern towards higher levels could then explain the size gradient observed within limpet populations.

This study shows that P. vulgata is well adapted to emersion, with higher respiration rates in air than in water, but that its calcification capacity limits the distribution of juveniles at the higher levels of the intertidal zone.


The paper

Tagliarolo M., Grall J. Chauvaud L. Clavier J., 2013. Aerial and underwater metabolism of Patella vulgata L.: comparison of three intertidal levels. Hydrobiologia 702:241–253.
See the first page


The authors

The four authors of this paper are members of the IUEM Laboratoire des sciences de l'environnement marin (Lemar)


The journal

Hydrobiologia was founded in 1948 and is published by Springer. This international journal is devoted to the biology of all aquatic environments and to the impact of human activities on them. It covers all organizational scales (molecule, organism, community, ecosystem) and includes disciplines such as limnology, oceanography, systematics and aquatic ecology.



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