Neohelice Research Articles

Freezing of movements and its correspondence with MLG1 neurons response to looming stimuli in the crab Neohelice.

Upon visually detecting a moving predator animals often freeze, i.e. stop moving, to minimize being uncovered and to gather detailed information of the object's movements and properties. In certain conditions the freezing behavior can be enough to avoid a predatory menace but, when the risk is high or increases to a higher level, animals switch strategy and engage in an escape response. The neural bases underlying escape responses to visual stimuli are extensively investigated both in vertebrates and arthropods. However, those involved in freezing behaviors are much less studied. Here, we investigated the freezing behavior displayed by the crab Neohelice granulata when confronted with a variety of looming stimuli simulating objects of distinct sizes approaching on a collision course at different speeds. The experiments were performed in a treadmill-like device. Animals engaged in exploratory walks respond to the looming stimulus with freezing followed by escaping. The analysis of the stimulus optical variables shows that regardless of the looming dynamic, the freezing decision is made when the angular size of the object increases by 1.4°. In vivo intracellular recording responses of Monostratified Lobula Giant Neurons (MLG1) to the same looming stimuli show that the freezing times correlate with the times predicted by a hypothetical spike counter of this neuron.

Effects of glyphosate on sperm quality of the estuarine crab Neohelice granulata, under the “One Health” perspective

This study was aimed at evaluating the effects of the commercial formulation Roundup Ultramax® on sperm mortality and viability, in terms of the capacity of spermatozoids (spz) to trigger the acrosome reaction (AR), using the estuarine crab Neohelice granulata as a model. To this, an in vivo assay comprising 100 days duration was carried out, on a control group and two groups exposed to the formulation (0.01 mg/L and 0.2 mg/L of glyphosate) under controlled conditions of photoperiod, feeding, and temperature. At the end of the assay, the right vas deferens (VD, proximal and middle portion) was dissected, and after homogenizing it in calcium-free saline solution, the acrosome reaction was induced in the phase containing the spz. In each treatment, the percentage of spz with total and partial AR was calculated, as well as that of dead spz. Compared to the control, crabs exposed to the herbicide showed a significant decrease in spz with full AR, together with an increase in the percentage of spz with partial AR. Furthermore, spz mortality was significantly higher in both glyphosate concentrations compared to the control, in a concentration-dependent manner. On the other hand, abnormal spermatophores, showing expanded walls and coalescence, were observed in a significant percentage in the left VD of the groups treated with the herbicide. The results obtained are compared with those from other studies on several invertebrate and vertebrate species that found inhibition of the AR and abnormal sperm, together with inhibition of spermatogenesis, endocrine disruption, and reduced sperm motility by effect of pure glyphosate and/or different glyphosate formulations. In summary, the available evidence highlights the possible impact of glyphosate on sperm quality, in a wide variety of species.

Noise accelerates embryonic development in a key crab species: Morphological and physiological carryover effects on early life stages

Anthropogenic noise is considered one important global pollutant. The impact of noise on marine invertebrates has been less assessed. The present study evaluated the chronic effect of the motorboat noise obtained from a lagoon's soundscape, the natural habitat of the key crab Neohelice granulata, on its whole embryonic development, considering morphological and physiological carryover effects on embryos and hatched larvae. Results demonstrated that embryonic development was shortened under noise exposure. The effects on advanced embryos, larvae and adult females were: increased heartbeats and non-viable eggs, and decreased fecundity. Biochemical responses showed lipid peroxidation in embryos while antioxidant enzymes were activated in larvae and adults, indicating a counteracting effect related to the life stage. The negative effects on fitness offspring may imply ecological consequences at the population level. Results are discussed in terms of the ecosystem engineer species studied and the habitat, a MAB UNESCO Reserve lagoon, suggesting the urgent need to develop mitigation plans.

Effects of high temperature and LPS injections on the hemocytes of the crab Neohelice granulata

Temperature fluctuations, particularly elevated temperatures, can significantly affect immune responses. These fluctuations can influence the immune system and alter its response to infection signals, such as lipopolysaccharide (LPS). Therefore, this study was designed to investigate how high temperatures and LPS injections collectively influence the immune system of the crab Neohelice granulata. Two groups were exposed to 20 °C (control) or 33 °C for four days. Subsequently, half were injected with 10 μL of physiological crustacean (PS), while the rest received 10 μL of LPS [0.1 mg.kg−1]. After 30 min, the hemolymph samples were collected. Hemocytes were then isolated and assessed for various parameters using flow cytometry, including cell integrity, DNA fragmentation, total hemocyte count (THC), differential hemocyte count (DHC), reactive oxygen species (ROS) level, lipid peroxidation (LPO), and phagocytosis. Results showed lower cell viability at 20 °C, with more DNA damage in the same LPS-injected animals. There was no significant difference in THC, but DHC indicated a decrease in hyaline cells (HC) at 20 °C following LPS administration. In granular cells (GC), an increase was observed after both PS and LPS were injected at the same temperature. In semi-granular cells (SGC), there was a decrease at 20 °C with the injection of LPS, while at a temperature of 33 °C, the SGC there was a decrease only in SGC injected with LPS. Crabs injected with PS and LPS at 20 °C exhibited higher levels of ROS in GC and SGC, while at 33 °C, the increase was observed only in GC and SGC cells injected with LPS. A significant increase in LPO was observed only in SGC cells injected with PS and LPS at 20 °C and 33 °C. Phagocytosis decreased in animals at 20 °C with both injections and exposed to 33 °C only in those injected with LPS. These results suggest that elevated temperatures induce changes in immune system parameters and attenuate the immune responses triggered by LPS.

Effect of low pH on embryonic and larval traits in the estuarine semi-terrestrial crab Neohelice granulata

The decrease in the pH of the oceans is one of the consequences of the release of CO2 (carbon dioxide) into the atmosphere by human activities. In estuaries, the magnitude to which pH will change and how the organisms that inhabit these environments would be affected is today relatively poorly known. We evaluated, by two laboratory experiments, whether the low pH affects early stages of the semi-terrestrial crab Neohelice granulata, an estuarine and ecosystem engineer species. Firstly, embryos and the spawning larvae were exposed to acidified seawater (ambient pH: 7.9 and 8, respectively; 7.5 and 6.9); secondly, ovigerous females were collected from the field and freshly hatched larvae were cultured under these pHs. We found a slight delay in the development time of embryos exposed to pH 6.9 but no difference in those reared under intermedial pH (7.5) with respect to the control and 6.9. We also found a reduction in egg volume and eye size exposed at pH 7.5 but with a recovery to normal size later in development. Heartbeats did not vary with pH. A delay in the duration of the intermolt period was observed for both larvae zoea I (ZI) and II (ZII) exposed to pH 6.9 in the first experiment, and only in ZI under this pH in the second one. Differences in body morphometry were found in the zoeas analyzed in the first experiment but not for the second. Measurements of changes in oxygen concentration conducted in the second experiment showed a reduction in the consumption of all zoeae stages when exposed to both low pHs. Mortality of larvae remained constant across treatments. Although it was found that the early stages of this species would be slightly/moderately affected by potential future scenarios of pH decrease, we cannot rule out major impacts at the population level, which requires further experiments to be carried out.

Insights into metal depuration in different tissues of the burrowing crab Neohelice granulata (Dana, 1852)

In this article, we address the limited understanding of the recovery abilities of estuarine organisms following exposure to metals and the lack of research on the mechanisms involved in metal sequestration and detoxification. Specifically, our focus is on the burrowing crab, Neohelice granulata, found in the Bahía Blanca estuary. The objectives of this study were to a) evaluate the metal content in various tissues of N. granulata before and after the depuration process, b) identify metal-rich granules (MRG) in the crabs' hepatopancreas and c) examine the relationship between metal loads and the induction/activity or reversibility of biochemical biomarkers. Following the depuration treatment, a noticeable decrease in metals was observed in the soft tissues compared to the hepatopancreas and the shell. Specifically, the depuration process effectively eliminated Mn and Fe in the form of MRG, but no significant differences were found for other metals, suggesting that MRG might not be the primary detoxification mechanism. Glutathione reductase (GR) and metallothionein (MT) levels significantly decreased after depuration. However, it is worth noting that the duration of the depuration period might not have been sufficient to fully counteract the oxidative stress and damage caused by field exposure.

Annual changes of Neohelice granulata cognitive abilities indicate opposition between short- and long-term memory retention

Neohelice is a long-standing model for memory studies for its strong retention of a reduced escape response when trained to iterative presentations of a visual danger stimulus (VDS). Here we present year-round changes that are related to the memory acquisition, storage, and expression. First, we evaluated exploratory activity and response to the VDS, as necessary for memory acquisition and expression. Both parameters change year-round. Second, short-term memory (STM) and two types of long-term memory (LTM) were assessed throughout the year. STM and long-term context-dependent signal memory (CSM) change between periods of the year, whereas signal memory (SM) does not, indicating that the cognitive abilities of the crab display circannual rhythms. Third, during the reproductive period, STM retention is higher than both CSM and SM, indicating a trade-off between STM and LTM. This is the first report of memory retention abilities changing seasonally as a trade-off between short- and long-term memories.

Unraveling the depuration mechanisms of metals in the burrowing crab (Neohelice granulata Dana, 1852): Biochemical biomarkers, metal-rich granules and bioaccumulation patterns

We studied the depuration mechanisms of metals (Cd, Cu, Pb, Zn, Mn, Ni, Cr, Fe) in Neohelice granulata, from sites with different human impacts (PC, a more impacted site and VM, a less impacted one). Our objectives included assessing metal concentrations (essential and non-essential) before and after depuration treatment, evaluating biochemical biomarkers (non-enzymatic and enzymatic) pre and post-treatment, and determining the role of metal-rich granules (MRG) in depuration. We observed variability in metals and biomarkers post-depuration, with no significant differences observed in PC, while Cd and Mn increased and Ni, Cu, and Fe decreased in VM. Integrated biomarkers' response indicated the prevailing antioxidant capacity in depurated organisms. Lipid peroxidation changes were insignificant, except in depurated-VM where values increased. MRG showed a significant decrease only for Mn and Fe, suggesting they were not the primary depuration structure. We concluded that depuration might depend on the species, gender and contamination history.

Discriminating the effects of bioturbation and herbivory by the intertidal burrowing crab Neohelice granulata on microphytobenthic assemblages of SW Atlantic mudflats

In shallow soft-bottom intertidal systems, microphytobenthic assemblages (MPB) and macrofauna interact and play important roles in ecosystem functioning. Macrofauna can affect MPB through bioturbation or/and herbivory. Most bioturbators also exert herbivory pressure on MPB, hence the results of the interaction between benthic macrofauna and MPB is not easy to predict. In this study we performed a small scale field experiment in order to discriminate the effect of bioturbation and herbivory by the intertidal burrowing crab Neohelice granulata on MPB structure, and primary production in SW Atlantic mudflats. The implemented treatments allowed us to separate the effects of bioturbation from those of herbivory on MPB. Results showed that N. granulata, through bioturbation + herbivory did not affect MPB structure (i.e., biomass, abundance and composition); and crab bioturbation without herbivory also did not modify MPB structure. Nevertheless, crab bioturbation without herbivory almost doubled MPB primary production, compared to the values registered in the treatment without bioturbation and herbivory and the treatment with bioturbation + herbivory. Sediment properties were equally affected by bioturbation in the four treatments. Thus, our results shows that the only significant effect on MPB structure is due to crab bioturbation.

Molecular insights from the crab Neohelice memory model.

Memory acquisition, formation and maintenance depend on synaptic post-translational machinery and regulation of gene expression triggered by several transduction pathways. In turns, these processes lead to stabilization of synaptic modifications in neurons in the activated circuits. In order to study the molecular mechanisms involved in acquisition and memory, we have taken advantage of the context-signal associative learning and, more recently, the place preference task, of the crab Neohelice granulata. In this model organism, we studied several molecular processes, including activation of extracellular signal-regulated kinase (ERK) and the nuclear factor kappa light chain enhancer of activated B cells (NF-κB) transcription factor, involvement of synaptic proteins such as NMDA receptors and neuroepigenetic regulation of gene expression. All these studies allowed description of key plasticity mechanisms involved in memory, including consolidation, reconsolidation and extinction. This article is aimed at review the most salient findings obtained over decades of research in this memory model.

Erratum: Eyestalk Ablation to Increase Ovarian Maturation in Mud Crabs.

Erratum: Eyestalk Ablation to Increase Ovarian Maturation in Mud Crabs.

Predatory behavior under monocular and binocular conditions in the semiterrestrial crab Neohelice granulata.

Neohelice granulata crabs live in mudflats where they prey upon smaller crabs. Predatory behavior can be elicited in the laboratory by a dummy moving at ground level in an artificial arena. Previous research found that crabs do not use apparent dummy size nor its retinal speed as a criterion to initiate attacks, relying instead on actual size and distance to the target. To estimate the distance to an object on the ground, Neohelice could rely on angular declination below the horizon or, since they are broad-fronted with eye stalks far apart, on stereopsis. Unlike other animals, binocular vision does not widen the visual field of crabs since they already cover 360° monocularly. There exist nonetheless areas of the eye with increased resolution. We tested how predatory responses towards the dummy changed when animals' vision was monocular (one eye occluded by opaque black paint) compared to binocular. Even though monocular crabs could still perform predatory behaviors, we found a steep reduction in the number of attacks. Predatory performance defined by the probability of completing the attacks and the success rate (the probability of making contact with the dummy once the attack was initiated) was impaired too. Monocular crabs tended to use frontal, ballistic jumps (lunge behavior) less, and the accuracy of those attacks was reduced. Monocular crabs used prey interception (moving toward the dummy while it approached the crab) more frequently, favoring attacks when the dummy was ipsilateral to the viewing eye. Instead, binocular crabs' responses were balanced in the right and left hemifields. Both groups mainly approached the dummy using the lateral field of view, securing speed of response. Although two eyes are not strictly necessary for eliciting predatory responses, binocularity is associated with more frequent and precise attacks.

First molecular identification of the trematode Maritrema bonaerense Etchegoin & Martorelli, 1997 (Plagiorchiida, Microphallidae) from its intermediate hosts, the gastropod Heleobia australis (d’Orbigny, 1835) (Littorinimorpha, Cochliopidae) and the crab Neohelice granulata (Dana, 1851) (Decapoda, Varunidae) in Argentina

The genus Maritrema Nicoll, 1907 (Platyhelminthes, Trematoda, Plagiorchiida, Microphallidae) comprises cosmopolitan species that predominantly parasitize birds. Although approximately 65 species have been described worldwide, including 6 for Argentina, molecular data referring to Maritrema species are still scarce worldwide, especially in South America. Unfortunately, this lack of references for nucleotide sequences is an obstacle to understanding the taxonomy and life cycles of trematodes, and impedes advancing our studies on the phylogeny and geographical distribution of these parasites. For that reason, we performed the molecular study of developmental stages of Maritrema bonaerense: cercariae (collected from the snail first intermediate host Heleobia australis, inhabiting Mar Chiquita lagoon) and metacercariae (collected from the crab second intermediate host Neohelice granulata, inhabiting Mar Chiquita lagoon and San Antonio Oeste, Argentina). The accordance between the ITS2 sequence of M. bonaerense cercaria from the snail H. australis and the sequences of metacercariae from the crab N. granulata was 100%, supporting previous findings of the life cycle of M. bonaerense based on morphological data. All Maritrema species are included in a monophyletic and well-supported clade. Maritrema bonaerense grouped more closely with Maritrema gratiosum. These findings contribute to the knowledge of digeneans in coastal marine ecosystems.

Assessment of metal contamination in an urbanized estuary (Atlantic Ocean) using crabs as biomonitors: A multiple biomarker approach

Estuarine environments, which are complex and sensitive coastal system, and are the final receptacle of several human wastes. Consequently, biomonitoring contaminants, such as metals, within these environments and developing scientific-based tools to conserve them have become particularly importat in recent years. Therefore, in the present study, we determined the levels of certain metals (Cd, Cu, Pb, Zn, Mn, Ni, Cr and Fe) both in sediments and in a key ecological benthic species, the burrowing crab Neohelice granulata, in sites of the Bahía Blanca estuary (SW Atlantic Ocean) with different anthropogenic impacts. We combined these records with geochemical indices and enzymatic and non-enzymatic biochemical biomarkers as early warning signals of contamination and damage to establish an integrated metal contamination profile. Our results indicated that some metals like Cu moderately contaminate the estuary and that the primary input of metals in this area is due to malfunctioning sewage discharges. Crabs exhibited the following pattern of metal accumulation: Fe ≥ Cu > Zn > Mn > Ni > Cd. Pb and Cr were under the method detection limit, and Cu was probably bioaccumulated from sediments. Metal concentrations showed significant differences according to the sites and seasons (p > 0.05) and not strictly to the crab gender. Besides, a similar pattern was observed for biomarkers, and the integrated biomarker response allowed establishing different oxidative stress patterns, according not only to human impacts but also to the seasonal physiological needs of this species and environmental endpoints (salinity, temperature, and pH). This work demonstrates that environmental factors also affect the metal influx in crabs and the activity of biomarkers beyond the source and fate of these elements. This information is vital for future integrated monitoring programs.

Tracking synthetic microdebris contamination in a highly urbanized estuary through crabs as sentinel species: An ecological trait-based approach.

Synthetic microdebris (particles of <5 mm) are a worldwide concern because they can affect the community structure of the aquatic ecosystems, organisms, and even food webs. For the biomonitoring of synthetic microdebris (especially microplastics, MPs), mainly benthic invertebrates are used, but crabs have been less studied in the literature. We studied the synthetic microdebris contamination in water, sediments, and three representative intertidal crabs (Neohelice granulata, Cyrtograpsus angulatus and Leptuca uruguayensis) with different lifestyles from the Bahía Blanca estuary, Argentina. The results obtained show the presence of cotton-polyamide (PA), polyethylene (PE), and polyethylene terephthalate (PET) in surface waters. In sediments, we identified cellulose modified (CE), polyester (PES), polyethylene (PE), and alkyd resin, while in crabs, cotton-PA and CE were the predominant ones. The MPs abundance ranged from 8 to 68 items L-1 in surface water, from 971 to 2840 items Kg-1 in sediments, and from 0 to 2.58 items g-1 ww for the three species of crabs. Besides, paint sheets ranged from 0 to 17 in the total samples, with Cr, Mo, Ti, Pb, Cu, Al, S, Ba and Fe on their surface. There were significant differences between the microdebris abundances in the abiotic matrices but not among crabs species. The ecological traits of the different crabs helped to understand the accumulation of synthetic microdebris, an important characteristic when determining the choice of a good biomonitor.

Matched function of the neuropil processing optic flow in flies and crabs: the lobula plate mediates optomotor responses in Neohelice granulata.

When an animal rotates (whether it is an arthropod, a fish, a bird or a human) a drift of the visual panorama occurs over its retina, termed optic flow. The image is stabilized by compensatory behaviours (driven by the movement of the eyes, head or the whole body depending on the animal) collectively termed optomotor responses. The dipteran lobula plate has been consistently linked with optic flow processing and the control of optomotor responses. Crabs have a neuropil similarly located and interconnected in the optic lobes, therefore referred to as a lobula plate too. Here we show that the crabs' lobula plate is required for normal optomotor responses since the response was lost or severely impaired in animals whose lobula plate had been lesioned. The effect was behaviour-specific, since avoidance responses to approaching visual stimuli were not affected. Crabs require simpler optic flow processing than flies (because they move slower and in two-dimensional instead of three-dimensional space), consequently their lobula plates are relatively smaller. Nonetheless, they perform the same essential role in the visual control of behaviour. Our findings add a fundamental piece to the current debate on the evolutionary relationship between the lobula plates of insects and crustaceans.

Inhibition of Oocyte Maturation in the Estuarine Crab Neohelice Granulata, by the Effect of Anti-Inflammatory Drugs.

The drugs ibuprofen and diclofenac were assessed in vivo on adult females of the estuarine crab Neohelice granulata. In a first, preliminary assay comprising 60-d, a significant (p < 0.05) lower content of total vitellogenic proteins was detected in the ovary at 10mg/L of each drug. In a second 90-d assay, comprising the exposure of crabs to 5mg/L of each drug during the entire pre-reproductive period, a significant (p < 0.05) decrease in the proportion of vitellogenic oocytes was observed by effect of diclofenac. The same effect was also observed in a third assay only comprising the last month of the pre-reproductive period, at 5mg/L of diclofenac, and also at a mixture of both drugs; besides, this mixture significantly (p < 0.05) increased the proportion of reabsorbed vitellogenic oocytes. The obtained results indicate that the effect of diclofenac is critical at the final stage of ovarian maturation, when the participation of prostaglandins is relevant.

Decapod egg membranes: powerful barriers or regulatory structures?

Osmoregulatory abilities and mechanisms of adults and larvae of decapod crustaceans have been extensively investigated. However, how embryos carried by their mothers can deal with changing or extreme salinities is less understood. The egg membranes are believed to isolate embryos from a challenging environment, although osmoregulatory ability has been demonstrated in early developing embryos (naupliar stage) of two crabs. To establish whether embryos are isolated by their membranes and/or are able to osmoregulate, we measured the survival and volume change over 48 h of oocytes and embryos in different stages of three carideans (Betaeus lilianae, Palaemon macrodactylus and P. argentinus) and the brachyuran Neohelice granulata, subjected to different salinities. In addition, we recorded osmolality changes in homogenates of the same stages in P. argentinus and N. granulata after 2 h of exposure and mapped the presence of putative sites of ions exchange in the membrane of all species. High mortality, when it occurred, was associated with low salinity and mortality variation with the stage of development depended on the species. All species precipitated silver salts in or under the egg envelope, with a different pattern between carideans and the brachyuran. Changes in osmolality and egg volume after hypo- or hyper-osmotic salinity challenges indicate that eggs are not fully isolated by their membranes, and that some osmoregulatory mechanisms are in play to maintain developmental homeostasis. We suggest that egg membranes can participate in osmoregulation by selectively transporting ions to an intramembrane space, with differences between carideans and brachyurans.

Contextual memory reactivation modulates Ca2+-activity network state in a mushroom body-like center of the crab N. granulata

High-order brain centers play key roles in sensory integration and cognition. In arthropods, much is known about the insect high-order centers that support associative memory processes, the mushroom bodies. The hypothesis that crustaceans possess structures equivalent to the mushroom bodies -traditionally called hemiellipsoid body- has been receiving neuroanatomical endorsement. The recent functional support is limited to the short term: in a structure of the true crab Neohelice granulata that has many insect-like mushroom bodies traits, the plastic learning changes express the context attribute of an associative memory trace. Here, we used in vivo calcium imaging to test whether neuronal activity in this structure is associated with memory reactivation in the long-term (i.e., 24 h after training). Long-term training effects were tested by presenting the training-context alone, a reminder known to trigger memory reconsolidation. We found similar spontaneous activity between trained and naïve animals. However, after training-context presentation, trained animals showed increased calcium events rate, suggesting that memory reactivation induced a change in the underlying physiological state of this center. Reflecting the change in the escape response observed in the paradigm, animals trained with a visual danger stimulus showed significantly lower calcium-evoked transients in the insect-like mushroom body. Protein synthesis inhibitor cycloheximide administered during consolidation prevented calcium mediated changes. Moreover, we found the presence of distinct calcium activity spatial patterns. Results suggest that intrinsic neurons of this crustacean mushroom body-like center are involved in contextual associative long-term memory processes.

Marine sources for biotechnology: Preliminary digestive enzymes assessment of seven decapod species from the southwest Atlantic

Marine natural compounds are required by industries to improve and reduce costs of many biotechnological processes. In this work, specific activity of midgut gland digestive enzymes was measured in seven species of decapods. Alkaline and acid peptidase, lipase and amylase activities were registered in Pleoticus muelleri (Pm), Artemesia longinaris (Al), Paralomis granulosa (Pg), Lithodes santolla (Ls), Munida gregaria (Mg), Neohelice granulata (Ng), and Ovalipes trimaculatus (Ot). The highest activities for all the analyzed digestive enzymes were obtained in Pg, Ls, Mg and Ot, which were used for optimum temperature and pH measurements and to test their stability at different temperatures and pH. In general, optimal enzyme temperatures were between 30 and 50 °C in the four crustaceans. Peptidases and lipases from these species showed in general alkaline optimal pH except for those from Ot which presented high activity at the whole pH range (2–12). Amylase from Ot presented its optimal at pH 6–8 and 30–50 °C. Thermal stability of peptidases was elevated at 30 °C for the four species tested (Pg; Ls; Mg and Ot), lipases were stable mainly at 30–50 °C and amylase was stable at 10 °C, 30 °C and 50 °C during the first 30 min of the assay. In general, stability of the four enzymes from Pg was greater at alkaline pH than at acid. The other species showed stability at all pH for the four enzymes. The range of temperature and pH of the tested enzymes activity are coincident with those used in various biotechnological processes and therefore indicates the potential of these enzymes to be used for industrial procedures. Further, most of the species studied are converted into fishery residues, either by processing or by-catch, so their reuse would also contribute to the sustainability of the activity.