Knapp, Jarred Lee (2015-12)
ENGLISH ABSTRACT: The West Coast rock lobster (WCRL), Jasus lalandii, is a critical marine fisheries resource for South Africa and may in future be negatively affected by the changes in seawater parameters associated with the ongoing anthropogenic carbon dioxide (CO2) emissions. These CO2 emissions have been linked to a global decrease in ocean pH (termed “ocean acidification”) and an increase in temperature. There are strong estimates that these changes are to worsen in coming centuries. This warranted research because of 1) the low current level of the resource (2.6% of pristine) and 2) the relatively unexplored physiological- and other biological responses of the WCRL to environmental stressors. This information is essential for the sustainable management of the resource by government scientists in times of global- and regional climate change.
In the short term, it was found that the WCRL was able to rapidly and reversibly respond to acute changes in seawater pH (pH 7.4), this was achieved primarily through the active up-regulation of bicarbonate levels in the haemolymph. Maintaining extracellular pH protects oxygen transport mechanisms, which are sensitive to pH changes due to the large Bohr effect that this study also revealed, in the respiratory protein, haemocyanin of adult WCRL.
The energy cost of actively maintaining extracellular pH, however, is expected to affect growth and potentially survival in the long term. This was tested on juvenile WCRL that were exposed to a reduced seawater pH of 7.3 (18.8 °C) over a period of 28 weeks. Results revealed that survival was not influenced and acid-base regulation in the hypercapnia-exposed lobsters was maintained throughout the duration of the trial, however, this led to a reduced growth rate. Subsequently, in order to replicate field conditions more closely, a combination of effects, namely seawater pCO2 (pH 8 and 7.3) and different temperatures (15.6 and 19 °C) on the growth of juvenile WCRL were assessed over an exposure period of 48 weeks in a second chronic trial. In contrast to the initial trial (28 weeks), where hypercapnia was assessed separately, lobsters exposed to hypercapnia had a higher growth rate than those at the same temperature exposed to a “natural” (normocapnic) seawater pH. The difference was interpreted as an indication that food availability/quality may negatively affect stress response, as feeding in the first trial was later considered “sub-optimal” in comparison to that of the second trial. In the latter, although both hypercapnia and temperature affected growth rates, temperature was the largest contributor to differences observed between treatments. The order of growth rates for lobsters from different treatments was: hypercapnia/high temperature > normocapnia/high temperature > hypercapnia/low temperature > normocapnia/low temperature. In this trial too, irrespective of treatment, lobsters were able to maintain extracellular pH within a relatively narrow range over the extent of the trial and survival was not negatively affected by hypercapnia or high temperature.
In order to compare the sensitivity of juvenile WCRL to that of adults, with regards to the effect of changes in extracellular pH on oxygen transport, and to assess the impact of chronic hypercapnia, haemocyanin from juveniles was studied in detail after the first growth trial. This revealed that juvenile WCRL have a similar Bohr effect to that of adults. In addition, the haemocyanin of hypercapnia-exposed juveniles showed an increased affinity to oxygen caused by an intrinsic change in its molecular structure. This was interpreted as an energy-saving mechanism, because at the same time, haemocyanin concentration in these animals was lower than in normocapnic lobsters.
At the termination of the second chronic trial, the immunological response to the combined stressors was assessed, namely total circulating haemocyte counts (THC) and the ability to clear/inactivate an introduced dose of a bacterium, Vibrio anguillarum. A pilot experiment on non-treated juveniles revealed a similar resting THC to that of other lobster species, and culturable V. anguillarum was rapidly cleared from their haemolymph. The effect of chronic exposure to a combination of effects, namely hypercapnia and different temperatures, was subsequently tested after termination of the second chronic trial. There were no differences between treatments in a) baseline THC (i.e. before bacterial challenge) and 2) the capability to clear culturable bacteria from haemolymph. The only difference was the circulating THCs post-bacterial challenge, as they were reduced in the hypercapnic-, high temperature treatment, compared with all other treatments. The reason is unknown, but it is speculated that it may have been linked to an increased metabolic demand in these lobsters.
Overall, these results demonstrate the great plasticity of the WCRL at the molecular-, biochemical and physiological level. They provide important initial information for government fisheries scientists to aid in predicting future development of, and potential threats to the WCRL resource, as well as providing a platform from which the direction of future studies can be determined.
AFRIKAANSE OPSOMMING: Die Weskus-seekreef, Jasus lalandii, is ’n belangrike seevisseryhulpbron vir Suid-Afrika en kan in die toekoms negatief geraak word deur die veranderinge in seewaterparameters wat met voortgesette antropogeniese vrystellings van koolstofdioksied (CO2) verband hou. Hierdie CO2-vrystellings word met ’n wêreldwye daling in die pH van seewater (oftewel “oseaanversuring”) en ’n temperatuurstyging verbind. Alles dui daarop dat hierdie veranderinge in die volgende eeue sal vererger. Dít regverdig navorsing weens 1) die huidige skaarste aan dié hulpbron (2,6% van oorspronklike getalle), en 2) die betreklik onverkende fisiologiese en ander biologiese reaksies van die kreef op omgewingstressors. Hierdie inligting is noodsaaklik om staatswetenskaplikes in staat te stel om die hulpbron te midde van wêreldwye en streeksklimaatsverandering volhoubaar te bestuur.
Op kort termyn word daar bevind dat die Weskus-kreef vinnig en omkeerbaar op akute veranderinge in die pH van seewater reageer (pH 7,4). Dít is hoofsaaklik deur die aktiewe opwaartse regulering van bikarbonaatvlakke in die hemolimf vasgestel. Die handhawing van ekstrasellulêre pH beskerm die meganismes wat suurstof vervoer, wat gevoelig is vir pH-veranderinge weens die beduidende Bohr-effek in die respiratoriese proteïen, hemosianien, by die volwasse kreef – nóg ’n bevinding van hierdie studie.
Tog sal die energiekoste verbonde aan die handhawing van ekstrasellulêre pH na verwagting groei en moontlik ook oorlewing op lang termyn beïnvloed. Dít is getoets op jong Weskus-krewe wat oor ’n tydperk van 28 weke aan seewater met ’n verlaagde pH van 7,3 (18,8 °C) blootgestel is. Resultate dui daarop dat oorlewing nié geraak word nie, en dat suur-basis-regulering in die hiperkapnie-blootgestelde krewe vir die volle duur van die proef gehandhaaf is, hoewel dit tot ’n verlaagde groeitempo gelei het. Ten einde natuurlike omstandighede akkurater na te boots, is ’n kombinasie van uitwerkings, naamlik pCO2 van seewater (pH 8 en 7,3) en verskillende temperature (15,6 en 19 °C), op die groei van jong krewe oor ’n blootstellingstydperk van 48 weke in ’n tweede chroniese proefneming beoordeel. In teenstelling met die aanvanklike proef (28 weke), is hiperkapnie afsonderlik beoordeel en het krewe wat aan hiperkapnie blootgestel is ’n hoër groeitempo getoon as dié by dieselfde temperatuur wat aan seewater met ’n ‘natuurlike’ (normokapniese) pH blootgestel is. Dié verskil is vertolk as ’n aanwyser dat voedselbeskikbaarheid/-gehalte ’n negatiewe uitwerking op stresreaksie kan hê, aangesien voeding in die eerste proefneming later as ‘suboptimaal’ beskou is vergeleke met dié van die tweede proef. In die tweede proef, hoewel hiperkapnie én temperatuur groeitempo’s beïnvloed het, was temperatuur die grootste bydraer tot die verskille wat tussen behandelings opgemerk is. Die orde van die kreefgroeitempo’s met die verskillende behandelings was: hiperkapnie/hoë temperatuur > normokapnie/hoë temperatuur > hiperkapnie/lae temperatuur > normokapnie/lae temperatuur. In die tweede proef kon die kreef ook, ongeag behandeling, ekstrasellulêre pH vir die volle duur van die proefneming binne ’n betreklik beperkte bestek handhaaf, en het nóg hiperkapnie nóg hoë temperatuur ’n negatiewe invloed op oorlewing gehad.
Om die gevoeligheid van jong Weskus-krewe met dié van volwasse krewe te vergelyk wat betref die uitwerking van veranderinge in ekstrasellulêre pH op suurstofvervoer, en om die impak van chroniese hiperkapnie te bepaal, is die hemosianien van jong krewe deeglik ná die eerste groeiproef bestudeer. Dít het aan die lig gebring dat die jong kreef ’n soortgelyke Bohr-effek as volwassenes toon. Daarbenewens toon die hemosianien van hiperkapnie-blootgestelde jong krewe ’n verhoogde affiniteit tot suurstof, wat deur ’n intrinsieke verandering in molekulêre struktuur veroorsaak word. Dít is as ’n energiebesparingsmeganisme vertolk, aangesien hemosianienkonsentrasie by hierdie diere terselfdertyd laer was as by normokapniese kreef.
Aan die einde van die tweede chroniese proefneming is die immunologiese reaksie op die gekombineerde stressors beoordeel, naamlik totale sirkulerende hemosiettellings (THC) en die vermoë om ’n toegediende dosis van die bakterie Vibrio anguillarum op te ruim/te deaktiveer. ’n Toetseksperiment met niebehandelde jong krewe dui op ’n soortgelyke rustende THC as dié van ander kreefspesies, en kweekbare V. anguillarum is vinnig uit die hemolimf opgeruim. Die effek van chroniese blootstelling aan ’n kombinasie van faktore, naamlik hiperkapnie en verskillende temperature, is vervolgens na afloop van die tweede chroniese proef getoets. Die verskillende behandelings lewer dieselfde a) THC op die basislyn (met ander woorde voor toediening van die bakterie), en 2) opruimingsvermoë van kweekbare bakterieë uit die hemolimf op. Die enigste verskil was die THC’s ná toediening van die bakterie, wat laer was met die hiperkapniese hoëtemperatuurbehandeling as met alle ander behandelings. Die rede hiervoor is onbekend, maar hou vermoedelik verband met ’n verhoogde metaboliese vereiste by hierdie krewe.
Oor die algemeen toon hierdie resultate die beduidende plastisiteit van die Weskus-seekreef op molekulêre, biochemiese en fisiologiese vlak. Dit bied belangrike aanvanklike inligting vir staatsvisserywetenskaplikes om die toekomstige ontwikkeling van én moontlike bedreigings vir die kreefhulpbron te voorspel, en voorsien boonop ’n platform van waar die rigting van toekomstige studies bepaal kan word.
Please refer to this item in SUNScholar by using the following persistent URL: http://hdl.handle.net/10019.1/97802