4 CDNICYT COMI5IÓ NACIONAL DE INVE5TIGAOÓN CIENTÍFICA Y TECNOLÓGICA
GOBIERNO DE CHILE
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COM1S1O NACIONAL DL INVESIJUAL ION CIENCIA Y 1 LCNOLOUI\
VERSION OFICIAL
FECHA: 16/3/2009
PROYECTO REGULAR N°1060127
INVESTIGADOR RESPONSABLE: THIEL MARTIN
FONDO NACIONAL DE DESARROLLO CIENTIFICO Y TECNOLOGICO (FONDECYT) Bernarda Morín 551, Providencia - casilla 297- y, Santiago 21 Telefono: 435 43 50 FAX 365 4435 Email: informes.fondecyt@ con¡ cyt.cI INFORME FINAL PROYECTO FONDECYT REGULAR RELACIONADO SR. MARTIN THI
N° PROYECTO: 1060127 DURACIÓN: 3 años AÑO ETAPA: 2008 TÍTULO PROYECTO: RAFTING DISPERSAL ON MACROALGAE DEPENDS QN FLOATINO TIME-FACTORS AFFECTING THE SURVI VAL OF FLOATING KELP AFTER DETACHMENT.
DISCIPLINA PRINCIPAL: BIOLOGIA MARINA GRUPO DE ESTUDIO: BIOLOGIA 1 INVESTIGADOR(A) RESPONSABLE: THIEL MARTIN DIRECCIÓN: Larrondo 1281, Casilla 117 RELACIONAD COMUNA: Coquimbo CIUDAD: Coquimbo SR. IVAN GOrv REGIÓN: IV REGION FONO: 5 1-209939 EMAIL: [email protected]
INFORME
OBJETIVOS
Cumplimiento de los Objetivos planteados en el Proyecto. Recuerde que los objetivos del proyecto no se refieren a listar actividades desarrolladas sino a los obietivos desarrollados N° OBJETIVOS CUMPLIMIENTO FUNDAMENTO 1 Examine the effect of water temperature and TOTAL grazing on fioatíng algae 2 Examine the effect of nutrients and temperature TOTAL on fioating algae 3 Examine the effect of PAR/UV-radiation and TOTAL - grazing 00 floating algae 4 Examine the effect of water temperature and TOTAL UV-radiation, on survival, growth and vitality of - floating Macrocystis spp. 5 Examine the effect of UV-radiation, on survival TOTAL growth and physiological performance of fioating Macrocystis spp. in their natural cnvironment.
Otro(s) aspecto(s) que Ud. considere importante(s) en la evaluación del cumplimiento de objetivos planteados en la propuesta original o en las modificaciones autorizadas por los Consejos.
In the original project, within the general objective to identif' the factors that influence survival and vitality of floating kelp, it was planned to do a long-term experiment over a time period of>2 months. However, experimental results from the tirst two years suggested that such an experiment will not produce new insights. Therefore, the objective 4 was modified to test thc combined effects of temperature and UVR, which was officially accepted by FONDECYT/CONICYT. Actually it was planned in the original project, to conduct objectivc 5 with the amphipod Peramphithoe femorata. However, during the summer 2008/09 this grazer was not very abundant in the kelp forests of the region of Coquimbo. We surveyed diftrent kelp forcsts nearby Coquimbo and tried to col lect them but we were unable to collect the large quantity of amphipods necessary for thc two cxperiments. For that reason and due to the fact that results from previous studies (objective 3) did not show any interactions between grazing and UVR, we finaily decided to concentrate our outdoor-laboratory and uield experiments only on the factor Uy. RESULTADOS OBTENIDOS: Please find attached in Appendix 1 figures" and in Appendix II "references'
OBJECTIVE 1 (Iaboratory : Examine the effect of water temperature and grazinc on fIoatinc algae.
OBJIa - Materials & Methods: In order to determine the effect of the factor temperature and herbivory on survival and vitality of floating Macrocystis spp. along the Chilean coast, an outdoor-tank experiment was conducted in Iquique (Universidad Arturo Prat), Coquimbo (Universidad Católica del Norte) and Calfuco (Laboratorio Costero de los Recursos Acuaticos, Universidad Austral de Chile) in austral summer 2007. Detached floating sporophytes were cultured at three different water temperatures and with or without the common amphipod grazer Peramphithoe femorata. Each treatment combination was replicated four times. At each location, we maintained floating algae at ambient temperature (incoming seawater), ambient temperature + 4t, and ambient temperature - 4'C (t ermed ambient, warm and coo!, respectively, in the following text). Whole sporophytes were placed in 90 L tanks and cultured for 14 days. Evaluations of growth parameters are briefly presented in the foliowing (for detailed results see Manuscripts 1 & 2).
OBJ la - Results and Interpretation: The results revealed that water temperatures between 12 1 and 20°C had only minor effects on the meristematic growth rates of Macrocystis spp., expressed as a daily blade elongation rate (BER) in cm d 1 (Fig. 1). However, water temperatures aboye 2M strongly influenced alga¡ degraciation and thus survival. At high water temperatures, floating individuals of Macrocystis spp. completely disintegrated and sank within 5 days (see Fig. 1, Iquique). Changes in algal biomass were dramatically different between the three locations. The alga¡ biomass (expressed as Percent biomass change d) considerably decreased at the warmest water temperature in Iquique (Fig. 2). In contrast, alga¡ biomass increased in Coquimbo and Valdivia. In Valdivia, alga¡ biomass continued to increase at ah temperatures (including the warm treatment) until day 10, even in the presence of grazers (Fig. 2). In Iquique, where algae were kept at warmer temperatures, algae continued to grow (albeit minimally - Fig. 1), but the biomass decreased dramatically under all experimental conditions (Fig. 2). In this case the high degradation rates could not be compensated by alga¡ growth. In Coquimbo alga¡ persistence mairily depends on the presence or absence of grazers. However, in Valdivia temperature throughout the tested temperature range and grazing seemed to have only minor impacts on alga¡ survival: algae grow and survive well due to very limited degradation rates. Physiological alga¡ responses were similar to growth responses of algae. Physiological performance of floating algae was favoured under optimal conditions such as low water temperatures and 10w hight levels, where algae were able to maintain a balance between the energy gained vía photosynthesis arid the energy used to cope with herbivores.The observed latitudinal persistence of floating Macrocystis spp. along the Chilean coast agrees with our previously predicted assumptioris that algae disintegrate at faster rates in warmer water temperatures (northern Chile) than algae kept at moderate and cooler water temperatures (central and south Chile). These results also agree with the findings of Vandendriessche et al. (2007) and Hobday (2000) that the decomposition rate of algae is temperature dependent. Using small-scale laboratory experiments, Vandendriessche et al. (2007) showed that alga¡ decomposition is minimal at 5 9C, and Hobd ay (2000) estimated that algal decomposition increase dramatically at temperatures abo ye 209C. Please find attached in the section products, subsection articles, Manuscript 1 & 2.
OBJECTIVE lb (field): Examine the effect of qrazinc on floatinQ al pae in the natural environment.
OBJ lb - Materials & Methods: In paraUel to the laboratory experiments, at ah three locations a field experiment was conducted. Three treatments were reahized, the Control (alga was kept in a mesh bag), Grazing (alga was kept in a mesh bag together with the amphipod grazer P. femorata) and Natural (alga was kept without a mesh bag). Each treatmerit was replicated 8-times. Algae were tethered in a protected bay and close to the location of the tank experiment to facilitate accessibility.
OBJ lb - Results: Prehiminary results show that the natural environment (temperature and herbivory) had shight effects on BER and biomass change. In Iquique the BER only seemed to be affected by the naturally occurring grazers in the field (Natural-treatment) and by P. femorata (Grazing-treatment, see Fig. 3). BER was strongly influenced by the amphipods in Coquimbo after a floating period of only 7 days and algae were so heavily grazed by the end of the experiment that it was not possible to take blade measurements. In Valdivia, during the first 7 days of experimentation BER was constant for al¡ treatments and decreased thereafter in the control and grazing treatment. Biomass decreased in the presence of P. femorata during the first 7 days of floating time in Iquique and Coquimbo, but not in Valdivia. Biomass change became negative between day 7 and 15 of floating time at ah three locations (Fig. 4). In general, there is a tendency that the mesh bags slightly suppressed algal growth when compared to the Natural treatment. OBJ lb —Interpretation: There is evidence that raft persistence in Iquique and Coquimbo is affected by the presence of amphipod grazers. It is known that vegetative blades of Macrocystis integrifolia showed 10w defensive responses against amphipod grazers (Pansch et al. 2008) and thus were probably preferentially consumed by Peramphithoe femorata (Manuscript 3). At both sites (northern and central Chile) the loss of biomass was higher than in Valdivia. Biomass losses in Valdivia can probably be compensated by alga¡ growth. However, for alI three locations algae kept under natural conditions may survive. In our experiments original grazers had been removed from the experimental algae from the Natural" treatment, which may not be entirely representative of naturally floating algae, which usually contain sorne grazers. Fish predators around natural rafts may, though, keep local grazer stocks low.
OBJECTIVE 2a (Iaboratory : Examine the effect of nutrients and temperature on floatinq algae.
OBJ 2a - Materials & Methods: In order to determine the effects of nutrient availability and temperature on growth, survival and vitality of floating Macrocystis spp., an outdoor-tank-experiment was conducted in autumn 2007. Detached floating sporophytes were cultured at three different water temperatures in flow- through seawater tanks, both at natural nutrient concentrations and in nutrient-enriched water. Nutrient levels in the nutrient-enriched treatment were enhanced by a standardized fertilizer (Osmocote 18% N, 6% P, 12% K). Small mesh bags filled with 50g of fertilizer were hung in the tanks. The same procedure was used for controls except that mesh bags contained inert glass beads. In order to confirrn that Osmocote was effective at increasing the nutrient concentrations in the surrounding waters, we sampled water around both control and Osmocote containing mesh bags. Results showed that our fertilization manipulation significantly increased the nutrients surrounding experimental algae when compared with controls. Both nutrient treatrnents were done at three different temperatures, ambient, warm and cool (see aboye). Whole sporophytes were placed in 90 L tanks and cultured for 20 days. Each treatrnent combination was replicated 4-times, i.e. 4 nutrient-enriched tanks and 4 ambient-nutrient tariks at each tested temperature. During and at the end of the experiment, samples for physiological and growth responses of algae were taken. Data of blade elongation rate and biomass change have also been recorded and are presented briefly in the following.
OBJ 2a - Results: At all ternperature and nutrient treatments tested, blade elongation rates (BER) of Macrocystis spp. (Hg. 5) decreased towards the end of the experiment. Lowest BER was recorded for the warm temperature treatment (18.8 ± 0.7t) independe nt of the presence of nutrients. Algae kept at the coolest water temperature (11.5 ± 1.1 1C) and in the presence of nutrient enrichment, however, showed highest persistence during the 20 days of experimentation. Between days 15 and 20 of the experiment, algae in the cool temperature and nutrient-enriched treatrnent showed highest growth rates compared to the other treatrnents tested. This suggests that 10w water temperatures in combination with high nutrient concentrations might enhance raft survival. AlI sporophytes gained in biornass until day 15 of the experirnent (Fig. 6). However, between days 15 and 20 of the experirnent, algae kept in the warm treatment Iost biomass independent of nutrient addition. In contrast, Macrocystis spp. cultured at cool (11.5 ± 1.1'C) and ambient (13.7 ± 1.2)ternperatures and in the ambient and nutrient enriched treatment gained in biornass until day 20.
OBJ 2a - Interpretation: There was little evidence that nutrient enrichment can increase growth performance of floating algae, which is in accordance with the results from Pfister and Van Alstyne (2003) who detected a slight increase in the growth of Hedophyllum sesslle after nutrient enrichrnent. However, the observed persistence of floating Macrocystis spp. agrees with our results from the ternperature experirnent (objective la): algae kept in warm water ternperatures showed lower BER and thus faster disintegration rates than algae kept at rnoderate and cool water temperatures. Results imply that raft survival in cooler water ternperatures can possibly be enhanced by sufficient nutrient availability in the surrounding water rnasses. In contrast, floating sporophytes travelling in warm waters do not have the capacity to increase their persistence with increased nutrient levels. Warm water temperatures seemed to have great negative impacts on alga¡ survival (Hobday 2000, Vandendriessche et al. 2007), independent from the addition of nutrients.
OBJECTIVE 2b (field): Examine the effect of nutrient availabilit y on Qrowth and vitalit y of floating Macrocystis spp in the natural environrnent.
This experiment was conducted parallel to the outdoor-tank experiment in auturnn 2007, where we wanted to estirnate the importance of nutrient availability in situ. Growth of kelps is mainly limited by nitrate (Brown et al. 1997). While attached algae gain sufficient nutrients from passing waters, floating algae move along within the same water rnass, and therefore may drive nutrient concentrations towards 10w values (Edgar 1987). Thus, thefloating potential of algae may be negatively affected by low nutrientievels. Sixteen sporophytes were individually tethered to a une in the Herradura Bay. One half of these sporophytes received small mesh bags filled with 50g of fertilizer (see abo ye) while the other half received mesh bags with inert glass beads. These mesh bags were changed every day in order to ensure that nutrients for algae were replenished every time. However, water samples that were taken around both treatments revealed no significant differences between control and nutrient enriched algae. Therefore, within this experiment, ah algae grew within the same nutrient conditions. In general it is difficult to experimentally enhance nutrients in the natural environment. However, we were able to successfully enhance the nutrients in a controlled laboratory experiment and results obtained allow relatively good estimations about the effect of nutrients on the floating potential of kelps.
OBJECTIVE 3 (Iaboratorv): Examine the effect of PAR/UV-radiation and qrazinq on floating akjae. OBJ 3 - Materials &Methods: In order to examine the effect of UV-radiation (termed UVR in the fohlowing) and herbivory on the growth and survival of floating Macrocystis spp., along the Chilean coast, three identical outdoor-tank-experiments were conducted in Iquique, Coquimbo and Calfuco. Experiments were carried out during austral summer in the time period between January and March 2008. Sporophytes were cultured in 90 L tanks under two different UV-regimes for which the experimental units were covered by standardized UV-filter foils. One foil blocked the UVR (Ultraphan 395, Digefra GmbH, Munich, Germany; letting only the photosynthetic active radiation PAR pass through), while the other foil ahlowed the complete transmissjon of ambient solar irradiance (PAR+UV, Ultraphan 295, Digefra GmbH, Munich, Germany) To test for filter artefacts, sporophytes in one set of tanks were cultured without filter foils (NATURAL). Additionally, sporophytes were cultured with or without the amphipod grazer Peramphithoe femorata. Four replicates were conducted for each treatment combination, i.e. 4 tanks with grazers and 4 tanks without grazers at each UV-treatment and NATURAL. Every day from 9:00 to 20:00 h, UVAB and PAR values were recorded using a LI-COR data logger. Maximum intensities of PAR and UVAB-radiation were recorded between 13:00 and 15:00 h for aPI three locations. Highest UVAB values were measured in Iquique. For Coquimbo and Calfuco UVAB values were almost in the same range and similarly high. Highest PAR values were recorded for Iquique and Coquimbo. Additionally, water temperature in the tanks was monitored at 9:00, 12:00, 15:00 and 18:00 h. Experiments lasted 15 days and algae were sampled at days 0, 5, 10 and 15. At each sampling day, growth and physiological responses of algae were measured.
OBJ 3 - Results: Detached sporophytes of Macrocystis spp. showed active BER (Fig. 7) ¡n alI U y- (PAR, PAR+UVAB and NATURAL) and grazing-treatments tested. In the northernmost location (Iquique), where highest UVR was detected, algae kept with PAR alone seemed to show higher BER. In Coquimbo, BER did not depend on the different UV-treatments. OnIy between days O and 5, there was a slight tendency of higher BER in the UV-blocked treatment. Here, BER seemed to depend on the presence of amphipod grazers. At the souttiernmost Iocation (Calfuco), which showed similar UVAB values as in Coquimbo, BER between days 0-5 were not affected by the U y- and grazing-treatments. Additionally control algae cultured without UVR showed highest BER between days 5 and 10 as in comparison to the other treatments. However between days 10 and 15, BER decreased in presence of grazers. Changes in algal biomass were dramatically different between the three Iocatíons (Fig. 8). In Iquique sporophytes kept without grazers and under NATURAL conditions lost biomass within the first sampling days. Between days 5 and 10 of the experiment, al¡ grazed sporophytes Iost biomass independent from the different UV-treatments. However, between days O and 5, highest biomass increase was detected for the UV-blocked sporophytes, independent of the presence of grazers. In Coquimbo al¡ control sporophytes gained in alga¡ biomass until day 15 of the experiment. Between days O and 5, algae treated without Uy (PAR) showed highest biomass increase. At day 5 grazed sporophytes had gained in biomass independent from UV-treatments, but Iost biomass during the following sampling intervals. OnIy between days 5 and 10, grazed UV-blocked algae increased their biomass. In Calfuco, alga¡ biomass increased until day 10 and even in the presence of grazers. Kelp physiology was only slightly affected by UVR in northern-central Chile and in southern Chile kelps could maintain their physiological performance.
OBJ 3 - lnterpretation: Solar radiation varied strongly along the Chilean coast. In the northernmost location, (Iquique) where highest UVAB-radiation was measured, the factor UVR together with grazing seemed to play an important role for the survival of floating algae. It is known that the growth of many benthic brown macroalgae is negatively affected by UVR, in particular during spring-summer season (Gomez et al. 2005; Roleda et al. 2006). In Coquimbo, alga¡ growth and thus persistence mainly depended on the presence or absence of grazers. In the southernmost location (Calfuco), where similar UVAB- radiation was detected as in Coquimbo, detached sporophytes showed a slight sensitivity to the factor UV and the factor grazing seemed to have only minor impacts on alga¡ survival. Similarly, Goméz et al. (2004) demonstrated for the same location that Macrocystis sporophytes are sensitive to the prevailing solar levels. Manuscripts on the effect of UVR and grazing on growth and physiological responses of floating kelps are in preparation and will be submitted to Marine Biology (since they have not yet been sent to the journal, they could not be uploaded on the CONICYT-website, but we can make them available upon request).
OBJECTIVE 4 (Iaboratorv): Examine the effect of water temerature and UV-radiation, on survival, growth and vitality of floatin q Macrocysfjs spp. In the original Fondecyt project, within the general objective to identify the factors that influence survival and vitality of floating kelp, it was planned to do a long-term experiment over a time period of >2 months. However, experimental results from the first two years suggested that such an experiment will not produce new insights. Therefore, this objective was modified to test the combined effects of temperature and UVR, which was officially accepted by FONDECYT/CONICYT.
OBJ 4 - Materials & Methods: In order to examine the effect of water temperature and UVR on growth and vitality of Macrocys fis spp., an outdoor-tank experiment was conducted, during austral summer 2008109 in Coquimbo. Detached sporophytes were cultured for 15 days in flow-through 90 L seawater tanks, at three different water temperatures similar as described abo ye (termed ambienf, warm and cao!) and under two different UV regimes. For (he two different UV-regimes the experimental units were covered by standardized UV-filter foils. One foil blocked the UVR (PAR), while the other foil allowed (he complete transmission of ambient solar irradiance (PAR+UV) (see also abo ye). Each treatment combination was replicated four times, i.e. 4 tanks that received only PAR and 4 tanks that received PAR+UV at each tested temperature. During experimentation, sporophytes were sampled at days 0, 5, 10 and 15. From al¡ sporophytes, samples were taken for enzyme, pigment and reproductive analysis. At this moment, analysis for pigments and enzymes are in progress as planned in the original project. Preliminary evaluations of blade elongation rate and biomass change have been conducted and will be briefly presented in the following.
OBJ 4 - Results: Preliminary results revealed that algae continuously elongated under al¡ water temperatures and Uy treatments tested (Fig. 9). BER decreased over time but independent of temperature and Uy treatments. Highest BER was detected at ambient water temperatures (17.1 ± 1.4 'C). Gain in alga¡ biomass was highest at water temperatures around 12 9C (Fig. 10). In contrast, at ambient water temperatures (17.1 ± 1.4 'C) biomass gains decreased after day 10 of the experiment in al¡ UV treatments. In the warm water treatment with temperatures> 19 'C, lowest biomass gain was detected an d biomass change was not affected by the presence (PAR+UV) and absence (PAR) of UVR.
OBJ 4 - Preliminary interpretation: It is known that UVR/PAR and elevated levels of temperature can negatively affect physiology and growth capacity of algae (e.g. Yakovleva et al. 2001, Lobban and Harrison 2000, Roleda et al. 2004). However, in our study, floating kelp seemed to be capable to adapt to their new light environment at the sea surface. Atiached individuals of Macrocystis integrifolla can react to UVA and UVB-radiation with the production of UV protective sunscreens such as phlorotannins (Swanson and Druehl 2002). Possibly, sporophytes of Macrocystis spp. in our study were able to produce these secondary metabolites while afloat. However, the factor temperature affected (he growth performance of floating Macrocystis spp. These results are in accordance with our previous results on temperature and alga¡ growth (see objective 1). While afloat, kelps are probably able to adapt to their changing light environment but the factor temperature plays an important role for the long distance dispersal of algae and its associated organisms.
OBJECTIVE 5a (field): Examine the effect of UV-radiation, on survival growth and physiological performance of floatin q Macrocysfis soo. in their natural environment.
In the original project it was planned to conduct objectives 5a & 5b with the amphipod Peramphithoe femorata. However, during the summer 2008/09 this grazer was not very abundant in the kelp forests of (he region of Coquimbo. We surveyed different kelp forests in (he vicinity of Coquimbo but we were unable to collect the large quantity of amphipods necessary for (he two experiments. For that reason and due (o the fact that results from previous studies (see objective 3) did not show any interactions between grazing and UV-radiation, we finally decided (o concentrate our outdoor-laboratory and field experiments on (he factor Uy. We conducted one field experiment (objective 5a) adjacent to the University in Bahía La Herradura, where artificial fioats were tethered. Moreo yer, we surveyed floating algae in (he bay system of Coquimbo along a south-north axis (objective 5b), which is representative of time as had previously been demonstrated in the thesis of Francisco Soto (see also "otros logros").
OBJ 5a - Materials & Methods: In arder (o estimate the importance of UVR on kelp floating in their natural habitat, kelp sporophytes were placed individually in small floats constructed with PVC tubes. Floats were covered by standardized UV-filter foils (with or without UV-radiation see abo ye). Each treatment combination was replicated 10-times, i.e. 10 sporophytes that were covered with PAR+UV filter foils, 10 sporophytes that were covered with PAR filter foils and 10 sporophytes (hat were kept without filter foils to test for eventual filter artifac(s (NATURAL). The experiment lasted 15 days and sporophytes were sampled at days 0, 5, 10 and 15. Additionally, irradiances were recorded from 9:00 to 18:00 h (local time). Besides (he growth measurements, samples for enzyrnes, pigmen(s and reproductive performance of algae were taken. This experiment was finished in February 2009, (herefore analysis of physiological alga] responses is still in progress and in the foliowing we briefly present growth performance of floating kelps. OBJ 5a - Results: Highest irradiances of PAR and UVAB were measured in the early afternoon around 15:00 h (local time), with PAR values of 1845.4 pmol m .2 -1, UVA values of 66.6W m 2 and UVB values of 2.4 W m -2 Preliminary results of BER showed a decrease of meristematic growth in time, with lowest growth activity at the end of the experiment (Fig. 11). Between days O and 5 of the experiment, blade elongation rafe was highest in the PAR and PAR+UV treatment, while a decline in BER was observed in the NATURAL treatment. Similarly, between days 5 and 10 of the experinient, BER was higher in the PAR and PAR+UV treatment than in the NATURAL treatment. Biomass change of floating algae considerably decreased in the PAR+UV and NATURAL treatment after day 5 of the experiment (Fig. 12). In contrast, floating sporophytes treated without UVR (Le. PAR treatment) gained in biomass until the end of the experiment. This suggests that UVR can have a damaging effect on alga¡ growth performance and that algae exposed (o the UVR aL the sea surface may experience lower survival times.
OBJ 5a - Preliminary ¡nterpretation: Herein, UVR negatively affected growth capacity of floating individuals of Macrocystis spp. These findings are in agreement wi(h the study by Clendennen et al. (1996), who showed that moderate U y leveis can have dramatic negative effects on the photosynthesis of Macrocystis pyrifera from southern California. Similarly, Macrocystis pyrifera from Mexico, which was cultured in large outdoor tanks, showed an increasing photoinhibition as a consequence of high exposure to UVB-radiation (Aguirre-von-Wobeser et al. 2000). A reduction in growth is the final direct consequence of these UV-mediated impacts on alga¡ physiology (Karsten et al. 2008). We suggest that kelps floating in regions with intense UVR may lose biomass faster than rafts floating in regions with low UVR. Probably, long distance dispersa¡ of kelps might thus be favoured under low UVR conditions, which depends e.g. on the cloud cover and the angle of the sun (i.e. the latitude).
OBJECTIVE 5b (fleid survey of floatinq alqae in a downstream gradient): Examine physiological performance and epibiont cover of freely floatinq Macrocy.s fis spp. in time.
OBJ 5b - Materials & Methods: In the bay system of Coquimbo, between Punta Lengua de Vaca (30114'S, 71`37'W) 29and Punta Choros (29014'S, 711 121W) (Fig. 13), freely floatíng algae were followed intime, in order (o determine their epibiont cover and (heir physiological performance. For this survey we exploited the fact (hat algae generaily move in northward direction in (his bay system (see thesis F Soto, section products, subsection tesis/memorias). Natural rafts were surveyed at six different stations within the bay system. Rafts were sampled with a dip net and brought to the laboratory. In total, eight plants from each station were used to conduct physiological measurements with a pulse amplitude modulation fluorometer (PAM-2000, Walz GmbH Germany). At the same time, from each of the eight alga¡ rafts, photographs from stipe, blade and pneumatocysts were taken in order to determine the percent epibiont cover by using Image- Pro plus 4.5. Epibiont cover was determined Lo estimate the approximate floating time of these natural alga¡ rafts. In addition, samples for enzyme and pigments were taken. Benthic algae from a kelp forest fo the south of (he bay system were used as controls. We herein present very preliminary data from three different stations aL the extremes of the north-south axis sampled along the bay system of Coquimbo.
OBJ 5b - Results and preliminary interpretation: Floating algae sampled at the 3 stations showed difterences in percent epibiont cover and in physiological responses (Fig. 14). There is a tendency that algae from the south with 10w epibiont cover (station 1 and 3) show higher physiological responses, than algae with high epibiont load (station 6). At station 1, epibiont cover is represented exclusively by diatoms, which suggests a very short floating time. However, at station 3 and 6 (he main colonizers are encrusting bryozoans, which colonized algae already after 5 days of detachment (in (he field experiment). Algae with high percent cover of epibionts are thought to be afloat for at least 20 days (station 6), whereas algae with 10w epibiont cover such as station 1 and 3 are estimated (o float since a few days (thesis F. Soto). Our results suggest that age of rafts (here estimated by epibiont cover) is negatively related (o (he physiological responses of floating algae, probably because of the floating time and also due (o the shading effects of epibionts. Algae can adjust their photosynthetic apparatus in response to variations in Iight environment and are classified in sun- or shade-adapted species, depending on their light requirements for the saturation of photosynthesis. Kelp floats with low epibiont cover and thus with low floating time, react like sun-adapted algae while the contrary is true for floats that show high epibiont cover. Low epibiont cover probably may favor algae in terms of protection against high light regimes. On (he other hand, large colonies of encrusting bryozoans can drastically reduce the amount of light reaching (he kelp surface (Hurd et al. 2000) as it is evident herein that these kelp react as shade adapted species. Therefore high epibiont load may reduce the energy available fo the underlying kelp tissue for C-fixation and other physiological processes and may probably provoke a fast sinking of alga¡ floa(s. Besides, during their journey, floating kelps may also be subject to changing water temperatures, which also can strongly influence kelp survival times. Overali, low and intermediate epibiont cover may act as a sunscreen against high irradiances, while high epibiont cover may accelerate (he physiological damage of algae and thus may limit the floating potential of kelp. DESTAQUE OTROS LOGROS DEL PROYECTO TALES COMO: - Estadías de investigación. - Formación de recursos humanos exceptuando tesistas ya informados. - Actividades de difusión y/o extensión en la temática del proyecto. - Cualquier otro logro no contemplado en los ítem anteriores y que Ud. quiera destacar. La extensión máxima de esta sección es de 1 página (letra tamaño 10, Anal o Verdana).
FORMACION DE RECURSOS HUMANOS: Similar as in the first two years, two assistants have participated in the project and gained experience in research projects. Eva Rothusler, the principal assistant, together with Osvaldo Cerda were responsible for the Iogistics of ah experiments conducted in Coquimbo. They designed and constructed the UV*Temperature outdoor laboratory experiment and the UV fleid experiment. The two assistants have participated in ah experiments, being responsible for collection of algae, and samphing during arid at the end of the experiments. Moreover, Osvaldo Cerda conducted his master thesis within this project.
As in the two first years, the principal assistant of the project, Eva Rothausler, has participated in al! experiments and has signed responsible for the data basis generated during the project, and for preliminary analyses. Eva has also learned Lo handie the UV data logger, to take the PAM-measurements and to process the data on photosynthetic activity of the algae. During a research stay at the University of Rostock (Alemania), Eva hearned to conduct CHN and Mannitol analyses and to interpret the resulting data. This information is of great value for the present project, because it contributes to a better understanding of the photosynthetic dynamics of the experimental algae.
The thesis project by Francisco Soto "Composición y dinámica de comunidades de epibiontes sesihes presentes en Macrocystis integrifolia en dos condiciones distintas: algas flotantes a ha deriva y fijas al sustrato" provides complementary information about the epibionts on fhoating ahgae, which accelerate the disintegration of these algae. The objectives of this thesis project focus on the epibiont load of floating algae in a south-north gradient within (he bay system of Coquimbo arid thus help understariding the dynamics of floating ahgae in this region. In addition, the thesis of F. Soto hehped to create estimations about the floating time based on growth rates of associated organisms such as Lepas spp. and M. ¡sabe/leona. His results from a descriptive samphing and a field experiment show that (i) fhoating algae found at farther distances from benthic source populations have higher loads of epibionts, and (u) that epibionts develop dense populations on floating algae while nearby benthic algae remain ahmost free of epibionts.
Osvaldo Cerda has studied the resource allocation of brown seaweeds under different environmental conditions to fulfilh (he requirements to obtain the MSc degree at the University Católica del Norte in Coquimbo. One part of his thesis was about the hypothetical compensatory growth of Macrocystis integrifolia against grazing by the nest-dwelhing amphipod Peramphithoe femorata (see also cooperation international). The results show that kehp can compensate grazer-induced losses at moderate grazing pressure. Compensation at attacked sites occurs at the cost of diminished growth at other within-sporophytes sites. This work enormously contributed to a better understanding of kelp-herbivore interactions in northern- central Chile. Compensation of tissue losses due to herbivory is a scarcely investigated topic in marine ecology, in rehation to studies about structurah and chemical defenses. RESUMEN: Describa en forma precisa y breve el tópico general del proyecto, sus metas y objetivos y los resultados alcanzados. Utilice un lenguaje apropiado para la comprensión del público no especialista en el tema. Esta información podrá ser difundida. La extensión máxima de esta sección es de 1 página (letra tamaño 10, Anal o Verdaria).
Many marine organisms are passively dispersed via floating items of biotic and abiotic origin. Especially for organisms with non-planktonic ¡¡fe cycles rafting on floating items is considered the most important mechanism for dispersa¡. However a successful dispersa¡ and colonization of new habitats, which results in an enrichment of local biodiversity, is only possible when floating items survive sufficiently long at the sea surface. Macroalgae with specialized floating structures (bladders or gas filled spaces) represent one of the most abundant and suitable rafting substrata in the world's oceans. In particular the dispersa¡ of small herbivorous invertebrates like isopods and amphipods is favored by floating macroalgae, because the algae provide both food and refuge. However the lifetime of algae on the sea surface is limited and this may impose restrictions on the dispersa¡ potential of organisms associated with them. Previous estimates on the survival periods of floating algae after detachment are highly inconsistent and vary between days and months. This may be primarily due to the fact that indirect measurements such as the growth determination of associated stalked barnacles and morphometric alga¡ characteristics have been used to obtain estimates. The uncertainty of these estimates makes it difficult to obtain reasonable estimates for the potential dispersa¡ distances of floating algae and their associated organisms. Herein we tested how water temperature, riutrient availability, UV-radiation (UVR), and herbivory affect the floating potential of Macrocystis spp along the Chilean Pacific coast. This kelp species is very abundant along the Chilean coast and shows a high floating potential due to gas-filled bladders. The effect of the aboye abiotic and biotic factors on floating algae were tested in different factor combinations in both field and outdoor laboratory experiments Results from outdoor laboratory experiments showed that the giant kelp Macrocystis spp. has highest survival times at water temperatures <15t (southern Chile) where raft degradation was slow anci grazer-induced biomass losses were compensated by continuing alga¡ growth. In contrast, in northern Chile high water temperatures (>20°C) provoked rapid degradation of Macrocystis spp. rafts, anci at the highest temperature treatment (-24°C) aH experimental algae died after the first 5 days of the experiment, irrespective of grazer treatment. At intermediate temperatures (15°C - 20°C) such as commonly found in central Chile, alga¡ survival depended on the presence of associated grazers. In absence of grazers alga¡ rafts gained in biomass while grazing caused considerable losses of algal biomass. Similar results were detected for the physiological reaction of floating algae. High water temperatures provoked a decrease in photosynthesis while at Iow water temperatures the physiological functioning of the photosynthetic apparatus could be maintained. The survival of floating Macrocystis spp, at 10w water temperatures can possibly be enhanced by nutrient availability in the surrounding water masses. However, algae in nutrient-enriched waters showed only a slight increase of growth. Floating sporophytes travelling in warm waters do not have the capacity to increase their persistence with increased nutrient levels. Warm waters negatively impacted survival of algae, irrespective of nutrient treatments. High UVR as measured at low latitudes (northern Chile) together with grazing led to a reduction in kelp growth rates and thus seemed to play an important role in the survival of floating algae. Contrarily, in central Chile, alga¡ growth and thus persistence mainly depended on the presence of grazers. In the southernmost location (Calfuco), where similar UVR was detected as in Coquimbo, detached sporophytes showed a slight sensitivity to the factor U y and the factor grazing had only minor impacts on alga¡ survival. However, the negative effects of UVR on algal performance did not reach the magnitude observed for high temperatures. In Coquimbo, where the crossed effect of UVR and temperature on the survival of floating kelp was tested, only the factor temperature affected the performance of algae. However, in a subsequent field experiment it was evident that UVR negatively affected biomass changes of floating kelp. Results showed that kelps floating in treatments with UVR lost biomass, while kelps maintained without UVR (PAR alone) gained in biomass. A parallel field survey showed that the photosynthetic response of kelp rafts floating for long time periods (determined based on epibiont sizes) was lower than that of kelps that had only floated for a few days. Overali, temperate kelp species that float after detachment seem to be equipped for long distance dispersa¡, but their survival at the sea surface is strongly affected by water temperatures and UVR. At low latitudes (northern Chile), high water temperatures and high UVR can provoke high biomass losses, thereby limiting the survival and possible dispersa¡ distances of kelp rafts. At mid latitudes (central Chile), the floating potential of kelps is strongly affected by associated herbivores, while at high latitudes the survival potential of algae at the sea surface is high, thereby allowing for long distance dispersa¡. The results of this study thus confirm that abiotic factors such as temperature and UVR determine the survival of kelp rafts at the sea surface, thereby also influencing the dispersal potential of associated organisms. COOPERACIÓN INTERNACIONAL
N° Proyecto: 7080193 Nombre Colaborador (a) Extranjero (a): KARSTEN ULF Afiliación Institucional Actual: UNIVERSITAT ROSTOCK Fechas de estadía Desde :14/11/2008 Hasta :28/11/2008
Describa las actividades realizadas y resultados obtenidos. Destaque su contribución al logro de los objetivos del proyecto. Si es pertinente, indique las publicaciones conjuntas generadas, haciendo referencia a lo informado en la etapa Productos. Agregue en la etapa anexos la información necesaria. International Cooperation Project 2008: general objective (compensatory growth): Examine the possible compensatory growth of Macrocystis spp. in the presence of Peramphithoe femorata.
Background information: In the International Cooperation Project from 2007 we had examined the reactions of Macrocystis spp. tu grazing and their effects on grazers. Those results had suggested that subapical blades of Macrocystis spp. increased their elongation rates when attacked by grazers. For the International Cooperation Project 2008 we had hypothesized that the observed high elongation rate is due to the compensatory growth capacity of Macrocystis spp. Several marine brown algae such as Fucus and Ascophllum produce chemical feeding deterrents when grazed by herbivores (Van Alstyne 1988, Rhode et al. 2004). 1-lowever, a previous study suggested that Macrocystis spp. lack grazer-promoted inducible defences (Macaya et al. 2005).
It is kriown that the considerable energy requirement for compensatory growth in brown algae can be supplied by internal storage carbohydrates such as mannitol and laminaran (Toyoki et al. 2008). These photosynthates can be used by algae for growth and survival wheu environmental conditions are unfavourable (e.g. Iow irradiance levels; Kremer 1984, Gomez & Wiencke 1998). The specific hypothesis tested in this project was: The presence of herbivorcs in subapical blades of Macrocystis integrifhlia promotes a cornpcnsatory growth at directly attacked blades or in zones with active growth such as apical blades or apical nieristenis
Prof. IJlf Karsten from the University of Rostock sígniftcantly contributed to the understanding of compensator ttrowth ni Macroc'vstis spp. by updating methodologies for thc quantification of carbohydrates (laminarian and mannitol) in macroalagc. His work was mainly related tu trajo the principal investigators (M. Thiel and F. Tala) and project assistants (O. Cerda and E. Rothusler) in modem enzymatic techniques to measure carbohydrate contents of apical and subapical blades of Macrocystis spp. that werc maintained in prcscnce and absence of the amphipod grazer Peramphithoe fernorata.
Specific 0W. compensatory growth: Detennination of carbohydrate content of grazed versus ungrazcd apical and suhapical blades.
OBJ A - Materials & Methods: A total of 24 sporophytes with their complete holdfasts were distributed individually ¡ti 90 1. plastic tanks. One half of the containers were kept with amphipods, while the other half was kept as grazer-free controls. On each grazed sporophvte, five amphipods were directly placed onto fivc subapical blades, where artificial nests were made by curling up thc terminal portion of each blade using a rubber band. Sporophytes were measurcd at days 2,4,6, 8, lO, 12 and 14 ¡ti order tu quantify growth rates of both apical and subapical blades from grazed and ungrazed individuals. Growth rates were determined using the hole-punch method by Parke (1948). A perforation aboye the meristematic zone on each apical and subapical blade was done and subsequent measurements at 2-day intervais were performed. Additionally, small tissue samples from the same blades were taken at day 14 fór the determínation of carbohydrates. After two weeks of experimentation we obtained growth rates and final laminaran content of apical and subapical blades from grazed and ungrazed sporophytes.
OBJ A Results: Amphipods caused significant biomass losses un subapical blades. However, thesc subapical blades maintained clevated growth rates similar to ungrazed subapical blades on the control sporophytes (see Fig. 2 in Manuscript 4). lo contrast, apical blades from grazed sporophytes exhihited lower growth rates than apical blades from ungrazed individuals. The results of carbohydrate quantifications revealed that grazed apical and subapical blades havc lower (albelt non sigriificant) mannitol contents than ungrazed apical and subapical blades. On the other hand, grazed apical blades exhibited lower laminaran content than ungrazed blades, while grazed subapical blades revealed higher laminaran concentrations than ungrazed laminae (see Fig. 4 in Manuscript 4).
OBJ A -- lnterpretation: Moderate grazing pressures, as reflected in our experiment, allowed giant kelp Macrocystis spp. to exhibit compensatory growth against grazing of the nest-dwelling amphipod Peramphithoe femorata. The continuous high growth rates of grazed subapical blades might be supported by an increased short-term turnover of mannitol in grazcd blades, while laminaran rcmains constant as long-term reserve supply. It is also probable that compounds other than mannitol and laminaran might increase their concentration in grazed subapical blades (e.g. amino acids like alanine or glutamic acid). Additionally, the compensatory response can be affected by other factors than grazers, e.g. seasonal variations in growth rates and nutrient availability. Further investigations considering a synergistic effect of environmental factors would be important to improve our understanding of the interaction between giant kelps and herbivores in fluctuating environments like the northern-central coast of Chile. In conclusion, compensatory growth by Macrocystis spp. against specialist herbivores can be achieved only at low and moderate grazing leveis, because it has been demonstrated that elevated grazing pressures might destroy almost the entire kelp sporophyte (Rothiusier et al. 2009). For more details, please see attached in subsection "articles" Manuscript 4.
References: Gómez, 1.. and Wiencke, C. 1998. Seasonal changes m C, N and major organic compounds and their significance to morpho-functional proccsses in the endemic Antarctic brown alga Ascoseira mirabilis. Polar Biology 19:115-24. Kremcr, B.P. 1984. Carbohydrate reserve and dark carbon fixation in the brown macroalga, Laminaria hyperborea. Journal of Plant Physiology 117: 233-42. Macaya, E.('., Rothilusler, E., Thicl, M., Molis, M. and Wahl, M. 2005. Induction of defenses and within-alga Yariation of palatability in two brown algae from the northem-central coast of Chile: Effects of rnesograzers and tJV radiation. Journal of Experimental Marine Biology and Ecology 325:214-227. Rohde, S., Molis, M. and WahI, M. 2004. Regulation of anti-herbivore defence by Fucus vesiculosus in response to various cues. Journal ofEcology 92:1011-1018. Rothiusler, E., Gómez, 1., Hinojosa, LA., Karsten, U., Tala, F. & Thiel. M. 2009. Effect of temperature and grazing un growth and reproduction of floating Macrocystis spp. (Phaephyceae) along a latitudinal gradient. Journal of Phycology 45: in press. Toyoki, 1., Kurashima, A. and Maegawa, M. 2008. Effect of seasonal changes in the photosynthates mannitol and laminaran un maturation of Ecklonia cava (Phaeophyceae, Laminariales) in Nishiki Bay, central Japan. Phycological Research 56:1-6. Van Alstyne, K.L. 1988. Herbivore grazing increases polyphenolic defenses in the intertidal brown alga Fucus distichus. Ecology 69:655-663.
PRODUCTOS
ARTÍCULOS Para trabajos en Prensa! Aceptados/Enviados adjunte copia de carta de aceptación o de recepción.
Autor (a)(es/as) : Rothaeusler, E.: Gómez, 1.; llniojosa, 1.: Karsten, U.: Tala, F.: Thiel, M. Nombre Completo de la Revista : Joumal of Phycology Título (Idioma original) : Effect of temperature and graLing un growth and reproduction of floating Indexación : ISI ISSN Año: Vol.:
Páginas Estado de la publicación a la fecha : En Prensa Otras Fuentes de financiamiento, si las hay: Fellowship frorn the German Academic Exchange Service (DAAD, August 2005-July 2007) to E. Rothaeusler
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N°: 2 Autor (a)(es/as) : Rothaeusler, E.; Gómez, 1.; Hinojosa, 1.; Karstcn, U.; Tala, F. Thiel, M. Nombre Completo de la Revista : Joumal of Phycology Título (Idioma original) : Physiological performance of floating giant kelp Macrocystis spp. (phaeophvceae): latitudinal variability in the effects of temperature and grazing Indexación : ISI ISSN: Año: Vol.:
Páginas Estado de la publicación a la fecha : Enviada Otras Fuentes de financiamiento, si las hay Fellowship of (he Gei-man Academic Exchange Service (DAAD, August2005-July 2007) to E. Rothaeusler. 1
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3 Autor (a)(es/as) : Pansch, C.; Gómez, 1.; Rotliusler, E.; Veliz, K.; Thiel, M. Nombre Completo de la Revista : Marine Biology Título (Idioma original) : Species-specific defense stratcgics of vegetative versus reproductive blades of úle Pacif'ic kelps Lessonia nigrescens and Macrocystis integrifolia Indexación : ISI 1SSN: Año: 2008 155 N°: Páginas: 51-62 Estado de la publicación a la fecha: Publicada Otras Fuentes de financiamiento, si las hay Fe]lowship from the Mercator Stiftung GmbH to C. Pansch.
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4 Autor (a)(es/as) : Cerda, O.; Karsten, U.; Rothiusler, E.; Tala, E.: ThieL, M. Nombre Completo de la Revista : Journal of Experimental Marine Biology and Ecology Título (Idioma original) : Compensatory growth of the kelp Macrocystis integrifolia (Phaeophyceae, Laminariales) against grazing of Peramphithoe femorata (Amphipoda, Ampithoidae) in northcrn-central Chile. Indexación : ISI ISSN: Año: Vol.:
Páginas Estado de la publicación a la fecha : Enviada Otras Fuentes de financiamiento, sitas hay
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OTRAS PUBLICACIONES
Sin información ingresada.
CONGRESOS
N°: Autor (a)(es/as) Rothaeus]er, E.: Tala, F.; Gomez, 1.; Thiel, M. Título (Idioma original) Effect of controlLed water temperature on the survival of tloating apical tips of Macrocystis integrifolia Nombre del Congreso XXVI Congreso de Ciencias del Mar País: CHILE Ciudad Iquique Fecha Inicio 22/05/2006 Fecha Término 26/05/2006 Nombre Publicación Año: Vol.:
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2 Autor (a)(es/as) : Rothaeusler, E.; Tala, F.; Gomez, 1.; Karsten, U.; Thiel, M. Título (idioma original) : Effekte vier verschiedener Meerwassertemperaturen auf das Überleben treibendcr apikaler Pflanzenteile der Braunalge Macrocystis integrifolia Nombre del Congreso : 11. Wissenschaftliche Tagung der Sektion Phykoiogic in der Deutschen Botanischen Gesellschaft País: ALEMANIA Ciudad : Helgoland Fecha Inicio : 28/08/2006 Fecha Término : 31/08/2006 Nombre Publicación Año: Vol.:
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N°: 3 Autor (a)(es/as) : Rothaeusler, E.; Tala, F.; Gomez, 1.; Hinojosa, 1.; Miranda, L; Karsten, EJ.; Thiel, M. Título (Idioma original) : lncreasing seawater temperatures limit dispersa] distances of kclp rafts Nombre del Congreso : 42. European Marine Biology Symposium País: ALEMANIA Ciudad : Kiel Fecha Inicio : 27/08/2007 Fecha Término: 31108/2007 Nombre Publicación Año: Vol.: N°: Páginas Envía documento en papel: no Archivo Asociado : EMBS2007_abstract_1060127.pdf htIp:/.cvalc l.conic yt.cI:ttiIorrne acadernteoindcx.php/investigadorrf4 congresos/descarga/14672797/ 10601275677
N°: 4 Autor (a)(es/as) Thiel, M.; Rothaeusler, E.; Gornez, 1.;Hinojosa, 1.; Karsten, U.; 1ala, F. Título (Idioma original) Rafting dispersal by floating kelp-stopped short by high seawater temperatures Nombre del Congreso 37. Benthic Ecology Meeting País: ESTADOS UNIDOS DE AMERICA Ciudad Rhode Island Fecha Inicio 09/04/2008 Fecha Término 13/04/2008 Nombre Publicación: Año: Vol.:
Páginas:
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5 Autor (a)(es/as) : Thiel et al. Título (Idioma original) : Viajes con origen y destino desconocido objetos flotantes y su rol ecologico en la costa Chilena Nombre del Congreso: XXVIII Congreso de Ciencias del Mar País: CHILE Ciudad : Viña del Mar Fecha Inicio : 26/05/2008 Fecha Término : 30/05/2008 Nombre Publicación Año: Vol.:
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6 Autor (a)(es/as) : Rothaeusler, E.: Gomez, 1.; Tala, F.: Hinojosa, 1.: Karsten, U.: Thiel, M. Título (Idioma original) : lncident solar ultraviolet radiation impair dispersa¡ distances of travelling kelp rafts Nombre del Congreso : 43. European Marine Biology Symposium País: PORTUGAL Ciudad : Ponta Delgada Fecha Inicio : 08/0912008 Fecha Término : 12/09/2008 Nombre Publicación Año Vol.: N' : Páginas: Envía documento en papel: no Archivo Asociado : EMBS2008_abstractj 0601 27.pdf http:/, 27/5689
TESISiMEMORIAS
Título de Tesis : Composición y dinamica de comunidades de epibiontes sesiles presentes en Macrocystis integrifolia en dos condiciones distintas: algas flotantes a la deriva y fijas al sustrato.
Nombre y Apellidos del(de la) Alumno(a) : Francisco Emiliano Soto Cortés Nombre y Apellidos del(de la) Tutor(a) : Martin Thiel Rubén Bulboa Título Grado: Pregrado Institución : Universidad Catolica del Norte País: CHILE Ciudad: Coquimbo Estado de Tesis: Terminada Fecha Inicio : 01/01/2008 Fecha Término : 30112/2008 Envía documento en papel : si Archivo Asociado : SOTOthesis 10601 27.pdf
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2 Título de Tesis : Resource allocation to defenses against herbivores and epibionts in brown algae from thc northern-central coast of Chile under different environmental conditions Nombre y Apellidos del(de la) Alumno(a) : Osvaldo Cerda Montenegro Nombre y Apellidos del(de la) Tutor(a): Martin Thiel Título Grado : Magister Institución : Universidad Catolica del Norte País: CHILE Ciudad: Coquimbo Estado de Tesis: En Ejecución Fecha Inicio : 01107/2008 Fecha Término: 31/03/2009 Envía documento en papel : si Archivo Asociado : CERDAthesis_l 0601 27.pdf
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ANEXOS
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N°: 3 Archivo Asociado : AcceptanceMS 1_1 0601 27.pdf http:/eva1cvt.conic ytclinfornieacadernicoindexphpinsestigadorf5 anexos dese argal 4672 797 10601 272529 4 Archivo Asociado : SubmissionMS2_1060127pdf lit tp:caIcyl.conicvt.ct/infornw acadernico/index.phpfinvestigador.f5 anexos/descargal 4677797/10601272S3(1
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