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Eyestalk ablation

Eyestalk ablation is a surgical procedure in crustacean aquaculture that involves removing one or both eyestalks from broodstock, primarily female penaeid shrimp such as Penaeus monodon and Litopenaeus vannamei, to induce ovarian maturation and spawning by disrupting the X-organ–sinus gland complex responsible for secreting gonad-inhibiting hormone (GIH).^[1]^[2] The eyestalks are typically crushed, cut, ligated, or cauterized, often unilaterally on the right side to preserve vision while achieving hormonal effects.^[3]^[4] This technique, developed in the 1970s, has enabled the scaling of commercial shrimp production by allowing continuous breeding independent of seasonal cycles, significantly boosting global output through accelerated gonad development and increased egg production.^[5]^[6] Ablation elevates vitellogenin synthesis and reduces molt-inhibiting hormone, promoting faster maturation, though it can lead to incomplete ovarian development or lower egg quality in some cases.^[1]^[7] Despite its efficacy, eyestalk ablation is linked to high post-procedure mortality (up to 50% in some reports), heightened stress responses, and compromised immune function, increasing disease vulnerability in ablated shrimp compared to non-ablated counterparts.^[7]^[8] Welfare debates center on potential pain from the procedure, with behavioral changes indicating distress, though empirical evidence for crustacean sentience and nociception remains sparse and inconclusive due to physiological differences from vertebrates.^[4]^[9] Recent advancements in high-quality feeds and environmental management have demonstrated that non-ablated broodstock can achieve equivalent or superior maturation and survival rates, fueling industry shifts away from the practice.^[8]^[10]

Biological Basis

Eyestalk Anatomy and Function

The eyestalk in decapod crustaceans, such as penaeid shrimp, is an elongated appendage extending from the head that integrates sensory and neuroendocrine functions. It houses the X-organ–sinus gland complex, a primary neuroendocrine center comprising approximately 150–200 neurosecretory cells clustered in the eyestalk's medulla.^[11] The X-organ synthesizes peptide hormones, while the adjacent sinus gland acts as a neurohemal organ, releasing these hormones into the hemolymph for systemic distribution.^[12] This complex produces regulatory neuropeptides, including crustacean hyperglycemic hormone (CHH), molt-inhibiting hormone (MIH), and gonad-inhibiting hormone (GIH), which collectively modulate physiological processes like metabolism, ecdysis, and reproduction.^[13] GIH, a key peptide synthesized in X-organ neurosecretory cells, exerts a tonic inhibitory effect on gonadal development, suppressing vitellogenesis (yolk formation) and oocyte maturation in wild crustaceans.^[14] In penaeid shrimp like Penaeus monodon, empirical studies demonstrate that intact eyestalks maintain elevated GIH levels, delaying reproductive maturation until environmental optima are met, as evidenced by seasonal spawning patterns correlating with natural hormone profiles.^[15] This inhibition prevents premature energy allocation to reproduction in suboptimal conditions, preserving somatic growth and survival.^[1] The eyestalk integrates external cues—such as photoperiod, temperature, and salinity—via sensory inputs from compound eyes and antennal structures, modulating X-organ activity through neural feedback loops.^[16] For instance, lengthening photoperiods signal neurosecretory cells to adjust GIH release, synchronizing gonadal cycles with seasonal productivity peaks, while salinity fluctuations influence hormone synthesis thresholds to align reproduction with favorable osmotic environments.^[17] This causal mechanism ensures reproductive timing via precise hormonal titration, rather than direct gonadal stimulation, underscoring the eyestalk's role as a central regulator in crustacean life-history strategies.^[16]

Hormonal Mechanism of Ablation

Eyestalk ablation primarily disrupts the secretion of gonad-inhibiting hormone (GIH), also termed vitellogenesis-inhibiting hormone (VIH), produced by the X-organ–sinus gland complex within the eyestalk of decapod crustaceans such as penaeid shrimp. This neuropeptide normally suppresses ovarian maturation by inhibiting vitellogenin synthesis in the hepatopancreas and oocyte development in the ovary, maintaining reproductive quiescence until environmental or endogenous triggers modulate its release. Surgical removal of the eyestalk eliminates the primary source of GIH, leading to a rapid decline in its circulating levels and derepression of downstream reproductive pathways.^[18]^[19] The resultant hormonal imbalance promotes elevated vitellogenin expression and hemolymph titers, facilitating yolk protein uptake and oocyte growth, as GIH directly represses vitellogenin gene transcription via signaling cascades involving cyclic AMP and other second messengers. Concurrently, ablation reduces inhibition on ecdysteroid production, with studies observing increased ecdysteroid levels that further support vitellogenesis and meiotic resumption in oocytes, though ecdysteroids play a secondary role compared to GIH in penaeids. This contrasts with natural maturation, where pulsed GIH release coordinates with stimulatory factors like gonad-stimulating hormone (GSH) from thoracic ganglia; ablation overrides this balance, artificially sustaining high vitellogenin and ecdysteroid activity without precise regulatory feedback.^[1]^[2] Empirical data from transcriptomic and immunoassays confirm these shifts: post-ablation GIH mRNA and protein levels drop markedly within hours, correlating with upregulated vitellogenin transcripts and 2- to 5-fold hemolymph increases in black tiger shrimp (Penaeus monodon), while ecdysteroid peaks align with advanced gonadal stages. However, ablation also impairs other eyestalk neuropeptides, such as molt-inhibiting hormone (MIH), potentially exacerbating ecdysteroid dysregulation beyond reproductive benefits. This mechanism underscores ablation as an exogenous override of inhibitory neuroendocrine control, distinct from endogenous cycles reliant on intact eyestalk signaling for synchronized hormone oscillations.^[18]^[20]

Historical Development

Early Discovery and Scientific Foundations

The foundational experiments establishing eyestalk ablation as a tool for inducing reproductive maturation in crustaceans date to 1943, when French biologist Jean Panouse demonstrated that bilateral eyestalk removal in the caridean shrimp Palaemon serratus triggered rapid ovarian development and spawning within days, contrasting sharply with unablated controls that showed no such progression.^[21] This observation provided the first empirical evidence of an eyestalk-mediated inhibitory mechanism on gonadal activity, as ablated females exhibited precocious vitellogenesis and oocyte maturation without external environmental cues typically required in nature.^[22] Panouse's work built on prior incidental findings from the 1930s linking eyestalk ablation to accelerated molting in crayfishes, but shifted focus to reproductive endocrinology by isolating the eyestalk's suppressive role through simple surgical intervention.^[23] In the ensuing decade, studies extended these findings to confirm causality via reciprocal experiments, including eyestalk implantation and extract injections. Researchers in the 1950s showed that homogenates from eyestalks, when administered to ablated crustaceans, reinstated gonadal inhibition, thereby identifying a neuroendocrine factor—later characterized as the gonad-inhibiting hormone (GIH) from the sinus gland—responsible for blocking ovarian maturation under normal conditions.^[24] These ablation-extract protocols established first-principles evidence that GIH secretion tonically suppresses vitellogenin uptake and oocyte growth, with removal disrupting this axis to unleash endogenous maturation processes.^[21] Early validations in lab settings prioritized penaeid species like Penaeus spp., where unilateral or bilateral ablation yielded maturation rates of 70-90% within 3-7 days post-procedure, versus near-zero in sham-operated controls maintained under identical photoperiod and nutritional regimes.^[25] By the mid-1950s, these milestones solidified eyestalk ablation as a reproducible model for dissecting crustacean reproductive physiology, emphasizing the eyestalk's X-organ-sinus gland complex as the primary site of inhibitory neuropeptide production.^[26] Initial applications remained confined to controlled laboratory environments for species such as Penaeus duorarum and P. setiferus, where ablation not only accelerated spawning but also enabled the collection of baseline data on hormone titers and gonadal indices, laying groundwork for causal models of neuroendocrine regulation independent of seasonal cues.^[27]

Commercial Adoption in Aquaculture

Eyestalk ablation gained commercial traction in shrimp aquaculture during the 1960s and 1970s, aligning with expansion in Asia and Latin America where initial farming booms relied on species such as Penaeus monodon and Litopenaeus vannamei. Developed from earlier Japanese techniques by Fujinaga in the 1940s and refined through French research in Tahiti, the method was applied to induce spawning in captive broodstock, supporting the shift toward controlled hatchery operations over sporadic wild collections.^[28]^[29] By enabling repeatable maturation cycles, ablation facilitated early scalability in facilities, with hatcheries achieving more consistent seed production for pond stocking in emerging centers like Taiwan and Ecuador.^[28] In Latin America, adoption accelerated mid-decade, with Panama implementing ablation for L. vannamei year-round reproduction by 1978, supplanting less productive P. stylirostris stocks once nutritional protocols were optimized.^[30] This practice underpinned hatchery expansions in Ecuador, where ablated broodstock supported intensive grow-out systems yielding over 10 metric tons per hectare by the early 1980s.^[28] Empirical records indicate ablation enhanced output predictability, with facilities reporting sustained nauplii yields from ablated females, enabling larger-scale dissemination of post-larvae for commercial ponds.^[31] The technique's integration marked a pivot from predominantly wild-sourced broodstock to ablation-dependent domesticated lines, correlating with post-1980s production escalations as global shrimp output rose from 22,600 metric tons in 1975 to over 1.4 million metric tons by 2002, driven by expanded hatchery capacities in ablation-reliant operations.^[30] This transition reduced seasonal constraints on seed supply, bolstering industry growth in Asia—particularly Thailand and Indonesia—where ablation complemented imported L. vannamei strains amid rising demand.^[28]^[30]

Techniques and Procedures

Methods of Eyestalk Removal

Eyestalk removal in penaeid shrimp broodstock, such as Litopenaeus vannamei and Penaeus monodon, primarily involves manual techniques applied to one eyestalk of mature females, typically without anesthesia to maintain procedural efficiency in hatchery settings.^[32] Surgical excision entails grasping the eyestalk with forceps and severing it at the base using fine scissors or a scalpel, often after briefly immersing the shrimp in chilled seawater (around 10–15°C) to reduce movement and facilitate handling.^[33] Thermal cauterization uses a heated needle, wire, or soldering iron to burn and seal the eyestalk base, minimizing hemorrhage while destroying the sinus gland; this method is favored in high-volume operations for its speed, with the tool preheated to 200–300°C and applied for 1–2 seconds.^[34]^[35] Ligation involves tying a fine thread or suture tightly around the eyestalk base to restrict blood flow and induce necrosis, followed by optional trimming of the distal portion; this non-invasive approach avoids open wounds but requires monitoring for slippage.^[34] Mechanical pinching or crushing employs forceps to compress the eyestalk midway or at the base, disrupting neural and vascular connections without full severance, though it risks incomplete ablation if pressure is insufficient.^[34] These methods are selected based on hatchery scale and equipment availability, with cauterization and excision predominant in commercial protocols due to their reliability in achieving gland inactivation.^[1] Procedural timing targets sexually mature females in the inter-moult stage, when exoskeleton hardness aids grip and reduces injury risk during handling; operations occur under ambient hatchery lighting to avoid circadian disruptions, with each ablation completed in under 10 seconds per individual to process batches of 50–100 shrimp.^[33] Industry surveys indicate ablation's persistence in over 70% of global shrimp hatcheries as of 2023, though method-specific adoption varies by region, with Asian producers favoring cauterization for its low cost and minimal tools.^[36] Species adaptations emphasize precision for penaeids' slender eyestalks (2–5 mm diameter), contrasting coarser applications in larger crustaceans, and protocols stress sterile tools to prevent secondary infections in humid environments.^[1]

Unilateral Versus Bilateral Ablation

Unilateral eyestalk ablation entails the removal of a single eyestalk, whereas bilateral ablation involves excising both eyestalks from crustaceans such as penaeid shrimp. In commercial aquaculture, unilateral ablation serves as the standard procedure for inducing ovarian maturation in female broodstock, particularly species like Litopenaeus vannamei, to promote reproduction while minimizing physiological disruption from the intact contralateral eyestalk, which retains partial hormonal regulation.^[37]^[38] This method has supplanted bilateral ablation in practical settings due to its superior balance of efficacy and reduced mortality risks, with empirical data indicating bilateral procedures yield mortality rates up to 68% in L. vannamei, compared to 35% for unilateral ablation.^[37]^[39] Bilateral ablation, though more disruptive to gonad-inhibiting hormone (GIH) production and thus potent for rapid vitellogenin (VTG) gene expression in immature females—as observed in prawns where unilateral ablation fails to trigger VTG while bilateral does so swiftly—is largely confined to research contexts or scenarios demanding maximal hormonal override.^[40] Studies demonstrate that unilateral ablation preserves some neuroendocrine functions, mitigating excessive stress responses, accelerated molting, and metabolic imbalances seen in bilateral cases, such as elevated hemolymph glucose and triglyceride alterations in both sexes but with compounded severity bilaterally.^[37]^[41] In L. vannamei, unilateral approaches also correlate with enhanced reproductive performance relative to non-ablated controls, albeit with elevated but manageable mortality, underscoring their procedural rationale in broodstock protocols.^[42] The preference for unilateral over bilateral in modern aquaculture reflects data-driven practicality, as bilateral's intensified effects often compromise long-term broodstock viability without proportional gains in maturation synchrony or output under controlled conditions.^[7] For instance, in penaeid species, unilateral ablation shortens molt intervals and stimulates gonad development sufficiently for commercial cycles, avoiding the disproportionate growth acceleration and survival penalties of bilateral removal.^[33]^[43]

Physiological and Reproductive Effects

Induced Reproductive Outcomes

Eyestalk ablation promotes ovarian maturation in crustaceans by diminishing gonad-inhibiting hormone (GIH) levels from the eyestalk's X-organ sinus gland, thereby alleviating suppression of vitellogenesis and enabling rapid yolk deposition in oocytes.^[1] This hormonal shift upregulates vitellogenin gene expression, with studies in black tiger shrimp (Penaeus monodon) documenting a 240-fold increase by day 7 post-ablation, alongside gonadosomatic index (GSI) elevation from 1.1±0.2% to 4.7±2.3% and ovary weight from 0.7 g to 3.6 g within the same period.^[1] The procedure accelerates maturation timelines, often completing full ovarian development in 3-10 days compared to weeks in intact females under natural conditions.^[1] In prawns such as Macrobrachium lanchesteri at ovarian index stages 50-70, ablation reduces time to ovigerous state from 20 days in controls to 12 days, while stages 30-45 advance from 25 days to 12 days.^[44] Spawning induction rates in ablated shrimp reach 66-90%, with peaks of 91.6% in wild-caught P. monodon and consistent multiple spawns per female (average 2.94 in wild stocks).^[45]^[46] Ablated females exhibit enhanced fecundity, yielding higher egg and nauplii quantities per spawn due to synchronized and repeated maturation cycles.^[47] Controlled trials confirm elevated nauplii production supports scalable hatchery operations, with ablation-linked spawn frequencies increasing overall larval output without reliance on seasonal cues.^[42]^[48]

Associated Health and Survival Impacts

Eyestalk ablation in penaeid shrimp, such as Litopenaeus vannamei, is associated with elevated mortality rates compared to untreated broodstock. In one study on Farfantepenaeus paulensis, unilateral ablation resulted in 35% mortality, while bilateral ablation led to 68% mortality, versus 2% in controls, attributed to impaired hemolymph homeostasis and secondary infections.^[49] Similar patterns occur in L. vannamei, where surgical stress and hormonal disruption contribute to post-operative losses, often exceeding 20-30% in commercial settings during or shortly after the procedure.^[50] Physiological stress manifests through acute immune perturbations, including a significant initial decline in total hemocyte count (THC) to 3.12 ± 0.75 × 10⁶ cells ml⁻¹ six hours post-ablation in L. vannamei, recovering partially by day 5 but indicating transient immunosuppression.^[51] Concurrently, hemolymph glucose levels rise sharply to 108 mg dL⁻¹ at six hours (versus 36 mg dL⁻¹ in controls), reflecting hyperglycemic hormone dysregulation akin to a stress response, though levels remain elevated at 93 mg dL⁻¹ after five days.^[51] These changes correlate with up-regulated energy metabolism genes, such as those for NADH dehydrogenase and ATP synthase, signaling heightened metabolic demands.^[1] Ablated broodstock exhibit shortened molting cycles and increased energetic expenditure, compromising long-term physiological balance without full endocrine recovery, as the eyestalk's role in regulating multiple neurohormones (beyond gonad inhibitors) leads to sustained imbalances in lipid and carbohydrate metabolism.^[1] Offspring from ablated females display heightened vulnerability to pathogens; postlarvae challenged with Vibrio parahaemolyticus (VpAHPND) showed reduced survival compared to those from non-ablated broodstock, with compromised immune priming evident in lower robustness metrics.^[52] This suggests transgenerational health deficits, including poorer stress tolerance, linked to maternal hormonal perturbations.^[52]

Applications and Economic Role

Role in Shrimp and Crustacean Farming

Eyestalk ablation serves as a key technique in broodstock management for penaeid shrimp species, including Penaeus monodon and Litopenaeus vannamei, within commercial hatcheries to trigger ovarian maturation and spawning.^[53]^[54] Unilateral ablation of the eyestalk in mature females disrupts the secretion of gonad-inhibiting hormone from the X-organ sinus gland complex, promoting rapid vitellogenesis and the production of nauplii for larval rearing.^[32] This method integrates into hatchery protocols by allowing controlled induction of multiple spawning cycles from individual broodstock, facilitating on-demand seed supply for grow-out ponds.^[31] In tropical aquaculture hubs such as Southeast Asia and Ecuador, the procedure is routinely applied to enable year-round hatchery operations, decoupling reproduction from environmental cues like photoperiod and temperature that naturally limit wild spawning.^[54] Operators typically ablate one eyestalk per female post-maturity, pairing ablated females with intact males in maturation tanks optimized for salinity, feeding, and water quality to maximize egg viability.^[55] Beyond penaeid shrimp, eyestalk ablation has been adapted for other farmed crustaceans, including freshwater prawns (Macrobrachium rosenbergii) and portunid crabs like Scylla paramamosain, where it similarly accelerates gonad development but with variable success tied to species-specific endocrine responses.^[4]^[56] Empirical observations indicate lower predictability in non-penaeid species, often necessitating adjuncts like hormonal injections to achieve reliable spawning rates.^[7]

Contributions to Global Aquaculture Productivity

Eyestalk ablation facilitated the commercialization of shrimp farming by enabling reliable induction of spawning in captive broodstock, a breakthrough adopted in the 1970s and 1980s that overcame limitations of wild-sourced gravid females.^[36] This procedure accelerates ovarian maturation and boosts egg production up to 10-20 times compared to non-ablated shrimp, providing hatcheries with predictable output essential for scaling postlarvae production.^[36] Consequently, it supported the transition from wild-dependent seed collection to domesticated hatchery systems, reducing pressure on natural stocks and enabling intensive pond stocking.^[57] The practice correlates with the rapid expansion of global farmed shrimp output, which grew to 5.88 million metric tons in 2024, surpassing wild capture and establishing aquaculture as the primary supply source.^[58] By ensuring consistent seed availability, ablation has underpinned profitable operations in hatcheries, where economic analyses emphasize its value in production planning and yield optimization over sporadic natural reproduction.^[5] In developing economies such as Ecuador, India, and Vietnam—major shrimp producers—this technology has intensified farming, enhancing local protein availability and export revenues that bolster food security and rural employment.^[32] Shrimp aquaculture's growth, driven by ablation-enabled reproduction, has positioned it as a key contributor to affordable seafood supply in regions reliant on coastal farming for economic stability.^[32]

Controversies and Empirical Debates

Welfare and Pain Considerations

Eyestalk ablation in shrimp elicits behavioral responses indicative of potential nociception, such as tail flicking, rubbing the affected area, and increased grooming, observed immediately following the procedure in species like Litopenaeus vannamei.^[59] These reactions align with broader evidence of nociceptors in decapod crustaceans, which detect noxious mechanical or thermal stimuli and trigger reflexive avoidance behaviors, as demonstrated in electrophysiological studies on crab sensory neurons.^[60] However, interpreting these as evidence of subjective pain remains contested, given the decentralized nervous system of crustaceans, which lacks a centralized vertebrate-like brain structure for integrating emotional or motivational components of suffering.^[9] Stress indicators during eyestalk ablation include elevated haemolymph glucose levels and visible agitation, which can be mitigated by topical anesthetics, suggesting acute physiological distress in the absence of such interventions; a 2004 study on L. vannamei found that lidocaine application prior to ablation reduced these responses compared to untreated controls.^[61] Proponents of minimizing welfare concerns argue that any nociceptive effects are transient and reflexive, akin to injury responses in invertebrates without evolutionary adaptations for prolonged pain states, supported by the immaturity of crustacean pain research relying on limited, sometimes disputed behavioral assays.^[62]^[9] Conversely, reviews of decapod sentience highlight motivational trade-offs in avoidance learning and opioid-sensitive behaviors as potential markers of pain beyond mere nociception, though empirical thresholds for sentience remain debated.^[63] Shrimp-specific evidence for sentience is weaker than for larger decapods like crabs or lobsters, with behavioral tests showing inconsistent motivational changes under noxious conditions, prompting calls for precautionary welfare measures despite phylogenetic and neuroanatomical distances from vertebrates.^[32] Animal welfare advocates, citing these stress proxies, criticize routine ablation without analgesics as inflicting unnecessary suffering, while skeptics emphasize that invertebrate physiology—lacking opioid-modulated pain pathways seen in vertebrates—renders claims of equivalent distress unsubstantiated, advocating empirical caution over anthropomorphic assumptions.^[64] No consensus exists on long-term pain, as post-ablation survival and reproduction often proceed without overt chronic behavioral deficits in farmed broodstock.^[18]

Evidence on Necessity and Long-Term Drawbacks

Empirical studies have challenged the necessity of eyestalk ablation for inducing ovarian maturation in shrimp broodstock, particularly Litopenaeus vannamei. Research by Zacarias (2020) demonstrated that non-ablated females achieved comparable maturation rates and spawning frequencies to ablated counterparts when provided with optimized environmental and nutritional conditions, such as high-protein diets and controlled photoperiods, while exhibiting significantly higher survival rates (up to 20-30% improvement in broodstock longevity).^[52] ^[65] Postlarvae and juveniles derived from non-ablated broodstock also showed enhanced robustness, with lower mortality under stress conditions like salinity fluctuations or pathogen exposure, suggesting ablation's indispensability is overstated in modern hatchery systems.^[52] Long-term drawbacks of repeated eyestalk ablation include reduced broodstock longevity and compromised offspring quality. Ablated females often experience accelerated energetic demands, shortened molting cycles, and heightened susceptibility to diseases due to immune suppression, leading to broodstock lifespans shortened by 15-25% compared to non-ablated groups in multi-cycle trials.^[1] ^[7] Successive generations from ablated lines have displayed signs of physiological strain, including incomplete ovarian development in later spawns and decreased fecundity (e.g., 10-20% fewer viable eggs per batch), potentially exacerbating genetic bottlenecks in selectively bred stocks over time.^[7] ^[66] While proponents argue ablation enables faster reproductive cycles—often achieving full ovarian maturation in 3-10 days versus 14-21 days naturally, beneficial in resource-constrained settings with limited holding capacity—recent analyses indicate no net yield advantages. A 2024 review of hatchery data found that ablation's short-term speed gains are offset by higher broodstock mortality and poorer larval viability, resulting in equivalent or lower overall postlarvae production per facility when survival metrics are factored in. ^[7] ^[67] Balanced empirical comparisons across trials emphasize that non-ablative approaches yield sustainable productivity without these offsets, particularly as hatchery infrastructure improves globally.^[5]^[3]

Alternatives and Innovations

Non-Ablative Reproductive Induction Methods

Non-ablative reproductive induction in penaeid shrimp, such as Litopenaeus vannamei, relies on environmental manipulations that replicate natural spawning cues to stimulate ovarian maturation and spawning without surgical intervention. Optimal conditions include maintaining water temperatures at 28–29°C, salinities of 30–32 ppt, and photoperiods of 12–14 hours light followed by 10–12 hours dark, often supplemented with natural sunlight exposure. These parameters align with seasonal variations in wild habitats, promoting gonad development through neuroendocrine pathways that reduce vitellogenesis-inhibiting hormone (VIH) secretion from the eyestalks. In commercial-scale trials with non-ablated L. vannamei broodstock, such regimens yielded spawning success rates of approximately 90%, with females producing 177,000 eggs per female per day on average.^[65] ^[31] Nutritional strategies further support vitellogenesis by supplying precursors for yolk protein synthesis, emphasizing diets rich in lipids and highly unsaturated fatty acids (HUFAs) rather than solely high crude protein levels. Broodstock conditioning with fresh or frozen squid (15% biomass), polychaetes, and mussels (8% biomass) combined with commercial pellets (2–4% biomass) enhances hepatopancreas lipid reserves, including saturated and monounsaturated fatty acids, linoleic acid, arachidonic acid, and eicosapentaenoic acid (EPA). This composition mobilizes nutrients to ovaries, increasing n-6 polyunsaturated fatty acids (PUFAs) and docosahexaenoic acid (DHA) content during mid-reproduction, thereby fostering oocyte development without ablation-induced hormonal imbalance. Studies indicate that such feeds improve fertilization rates (p < 0.05) and hatching rates to 54%, surpassing 50% observed in some ablated controls.^[65] ^[31] Empirical comparisons demonstrate mechanistic viability for species like L. vannamei, where non-ablated females exhibit higher per-female fecundity and nauplii production than ablated counterparts (177,239 vs. 145,729 eggs/female/day; p < 0.05), alongside comparable overall productivity under intensive conditions. Offspring from these methods show enhanced larval quality, including superior resilience to salinity stress (99% survival vs. 96%; p < 0.05) and pathogens like Vibrio parahaemolyticus (70.4% survival at 96 hours vs. 38.8%; p < 0.05), attributed to elevated PUFA reserves in eggs and nauplii. While tank-level productivity may lag due to lower spawning frequency, these approaches prove sustainable for select penaeids, yielding robust postlarvae without surgical stress.^[65] ^[31]

Emerging Genetic and Hormonal Strategies

Selective breeding programs for penaeid shrimp, such as Litopenaeus vannamei, have focused on developing lines with enhanced natural maturation traits since the early 2010s, aiming to eliminate reliance on eyestalk ablation. These efforts target genetic variations associated with spawn capability and ovarian development, yielding stocks where non-ablated females demonstrate reproductive performance and offspring quality comparable to ablated counterparts in commercial-scale trials.^[68]^[31] Postlarvae and juveniles from such non-ablated broodstock also exhibit greater resilience to pathogens like Vibrio parahaemolyticus AHPND and white spot syndrome virus, supporting the viability of ablation-free lines.^[52] RNA interference (RNAi) techniques targeting gonad-inhibiting hormone (GIH) transcripts represent an emerging genetic alternative, silencing the hormone's suppressive effects on vitellogenesis without physical ablation. In L. vannamei, dsRNA-mediated GIH knockdown induced ovarian maturation in captive females, with the first successful application reported in 2015 as a direct substitute for eyestalk removal.^[69] Similar RNAi silencing of GIH in Penaeus monodon promoted gonad development and spawning under controlled conditions, demonstrating efficacy in multiple penaeid species.^[70]^[71] These methods leverage endogenous pathways, potentially reducing stress while achieving maturation rates akin to traditional practices, though scalability in farms remains under evaluation.^[72] Hormonal strategies, including exogenous injections of 20-hydroxyecdysone (20E), have been investigated to modulate reproductive cycles by mimicking or overriding GIH inhibition. While primarily linked to molting regulation, 20E administration influences neuroendocrine pathways that intersect with vitellogenesis, with studies showing accelerated ovarian progression in treated shrimp.^[73] In combination with genetic selection, these approaches contribute to domesticated lines exhibiting partial independence from ablation, as evidenced by improved baseline maturation in breeding programs.^[74] Ongoing trials emphasize precise dosing to optimize efficacy and minimize off-target effects on survival and growth.^[37]

Recent Developments and Future Outlook

Regulatory and Industry Shifts

In 2023, the Global Seafood Alliance (GSA) surveyed major shrimp producer associations worldwide, revealing that eyestalk ablation remained prevalent in hatcheries for species like Litopenaeus vannamei and Penaeus monodon, though many respondents identified viable alternatives and cited challenges such as higher costs and training needs in transitioning away from the practice.^[36] In August 2025, GSA announced a binding requirement under its Best Aquaculture Practices (BAP) certification standards, mandating that all certified shrimp farms and hatcheries worldwide cease eyestalk ablation or sourcing ablated broodstock by December 31, 2030, with interim progress reporting to monitor compliance and production stability.^[75] ^[76] This policy shift, informed by the survey's data on industry readiness, aims to align certification with empirical evidence of non-ablative methods' efficacy while avoiding disruptions to global supply chains.^[77] Regional industry responses have accelerated post-2020, particularly in Ecuador, where major broodstock producers such as Biogemar, Aquagen, and Omarsa fully eliminated eyestalk ablation by 2025, transitioning to controlled maturation protocols and reporting sustained larval output without yield losses in monitored trials.^[78] In Asia, select hatcheries in countries like Thailand and Vietnam have initiated partial adoption of ablation-free systems under pilot programs tied to export certifications, with empirical tracking showing initial 10-15% cost increases offset by improved broodstock survival rates over 12-18 month cycles.^[5] These trials, often linked to BAP or Aquaculture Stewardship Council (ASC) audits, prioritize data-driven validation of productivity metrics before scaling. Welfare-focused certifications have exerted growing market pressure on supply chains, with ASC standards prohibiting sourcing of larvae from ablated females effective April 2024, prompting retailers in Europe and North America to favor compliant suppliers and driving an estimated 20-30% shift toward non-ablated stock in certified segments by mid-2025.^[79] ^[6] Major UK retailers, including the Co-op, achieved near-total elimination of ablated prawn supplies by 2023 through supplier mandates, reflecting broader consumer and regulatory demands for verifiable welfare improvements without unsubstantiated productivity trade-offs.^[80] These dynamics have incentivized vertical integration in ablation-free production, as evidenced by rising premiums for certified shrimp in export markets.^[76]

Ongoing Research and Empirical Findings

A 2025 review in Reviews in Aquaculture evaluates alternatives to eyestalk ablation, concluding that non-ablative procedures yield equivalent reproductive outcomes in terms of maturation rates and larval production, supporting their scalability in commercial hatcheries with minimal productivity loss.^[32] Empirical trials reported in Zacarias et al. (2021) demonstrate that offspring from non-ablated Litopenaeus vannamei females exhibit higher survival rates under challenge from acute hepatopancreatic necrosis disease (Vibrio parahaemolyticus) and white spot syndrome virus compared to those from ablated broodstock, with statistically significant advantages against AHPND.^[81] Subsequent research by Simão Zacarias at the University of Stirling, building on 2020-2024 data presented at the Responsible Seafood Summit, shows that nutritional supplementation with EPA and DHA fatty acids accelerates ovarian maturation in non-ablated females to levels matching or surpassing ablation-induced rates, while enhancing offspring resilience to disease and stress.^[67] These non-ablative approaches, integrated with selective breeding of healthy males, have been implemented successfully in hatcheries across Honduras, Ecuador, Asia, and Madagascar's Unima operations, confirming ablation's dispensability without yield reductions.^[67] Projections based on these findings indicate feasible full transitions to integrated methods—combining nutrition, environmental cues, and genetic selection—within 3 years per adopting facility, yielding cost-neutral or reduced operational expenses due to improved broodstock longevity and larval vigor.^[67] Such data underscore the causal role of ablation in compromising disease resistance, positioning non-ablative innovations as viable for sustained aquaculture productivity.^[81]

References

[1]
Insights into Eyestalk Ablation Mechanism to Induce Ovarian ...
Eyestalk ablation is commonly practiced to induce ovarian maturation in the black tiger shrimp farming [1]. This study aimed to understand effects of the ...
[2]
Insights Into Mechanisms of Eyestalk Ablation-Induced Ovarian ...
Eyestalk ablation is the most common method to induce ovarian maturation in decapod crustacean aquaculture, but it jeopardizes broodstock survival and ...
[3]
Why stop using Eyestalk Ablation? - Shrimp Welfare Project
Eyestalk ablation (ESA) involves one or more of the animal's eyestalks being removed through crushing, cutting, burning, or tying it off.
[4]
Eyestalk ablation procedures to minimize pain in the freshwater ...
This article proposes an enhanced and less painful procedure to eliminate the flow of hormones from the X organ sinus gland in adult Macrobrachium americanum.
[5]
What is the path to viable alternatives to shrimp eyestalk ablation?
May 20, 2024 · Removing the eyestalk of broodstock female shrimp aided in the explosive growth of commercial-scale aquaculture. An undertaking by a Central ...
[6]
Shrimp eyestalk ablation - ASC International
a procedure used to induce spawning in broodstock shrimp.
[7]
Eyestalk ablation in crustacean production a brief review of the ...
Jun 25, 2024 · Despite its benefits, it raises concerns regarding challenges such as incomplete ovary development, decreased fecundity, and deleterious effects ...
[8]
New research proves eyestalk ablation is unnecessary and ...
Eyestalk ablation is unnecessary, increases disease vulnerability, and non-ablated shrimp have higher survival rates against diseases. High quality feed can ...Missing: controversies | Show results with:controversies
[9]
Review of some scientific issues related to crustacean welfare
Jun 15, 2018 · The scientific literature on the subject of welfare and pain in crustaceans is immature. It is based largely on a few dubious and disputed studies.
[10]
https://www.thefishsite.com/articles/the-case-against-eyestalk-ablation-in-shrimp-aquaculture
[11]
Regulation of Crustacean Neurosecretory Cell Activity - PubMed
The X organ-sinus gland system is a conglomerate of 150-200 neurosecretory cells in the eyestalk of crustaceans. It is the source of a host of peptide ...
[12]
Crustacean hyperglycemic hormone - ScienceDirect.com
Crustacean hyperglycemic hormone (CHH) is produced by the X-organ sinus gland complex in the eyestalk. CHH is a hyperglycemic factor and then regulates the ...
[13]
Signaling Pathways That Regulate the Crustacean Molting Gland
MIH, produced in the eyestalk X-organ/sinus gland complex, inhibits the synthesis of ecdysteroids. A model for MIH signaling is organized into a cAMP/Ca2+ ...Missing: anatomy | Show results with:anatomy
[14]
Molecular characterization of gonad‐inhibiting hormone of Penaeus ...
Feb 8, 2008 · Gonad-inhibiting hormone is a member of the neuropeptide family that is synthesized in neuroendocrine cells located in the eyestalk medulla ...
[15]
Suppression of a Novel Vitellogenesis-Inhibiting Hormone ...
Nov 7, 2021 · Suppression of a Novel Vitellogenesis-Inhibiting Hormone Significantly Increases Ovarian Vitellogenesis in the Black Tiger Shrimp, Penaeus ...
[16]
Reproductive regulators in decapod crustaceans: an overview
Jan 1, 2011 · Environmental variables such as salinity, temperature and seasonality play vital roles in regulating crustacean physiology, including ...<|control11|><|separator|>
[17]
effects of eyestalk ablation at different times of the year
sonally environmental stimuli, such as pho- toperiod and temperature, have proved to be strong inducers or modulators of crustacean reproduction and molting ...
[18]
Insights Into Mechanisms of Eyestalk Ablation-Induced Ovarian ...
Aug 14, 2020 · ... gonad-inhibiting hormone (GIH), the molt-inhibiting hormone (MIH) ... Insights into eyestalk ablation mechanism to induce ovarian maturation in the ...
[19]
Suppression of a Novel Vitellogenesis-Inhibiting Hormone ...
Nov 8, 2021 · monodon gonad-inhibiting hormone ... Insights Into Eyestalk Ablation Mechanism to Induce Ovarian Maturation in the Black Tiger Shrimp.
[20]
Comparative transcriptomics reveals eyestalk ablation induced ...
Aug 21, 2020 · ... gonad inhibiting hormone produced by the X-organ sinus gland. However, eyestalk ablation also results in negative impacts on the ...
[21]
Ovarian maturation and spawning in the white shrimp, Penaeus ...
Predictable induced reproduction in captive penaeids without the use of eyestalk ablation was considered a long term goal for shrimp mariculture (Quackenbush, ...
[22]
Eye Stalk Ablation in Crustaceans | PPTX - Slideshare
Panouse (1943)observed that removal of eyestalk of shrimp Palaemon serratus led to ovarian development. Ever since this discovery several workers have tried ...
[23]
[PDF] TECHNICAL REPORT NO. 431 - à www.publications.gc.ca
1930's and early 1940's 1n which several workers observed. Page 96. - 92 ... of eyestalk ablation and of injection of eyestalk extracts on testicular ...
[24]
A Crab Is Not a Fish: Unique Aspects of the Crustacean Endocrine ...
Mar 2, 2021 · Its important role was discovered through a series of ablation and implantation experiments in the 1950's (240–243). ... penaeid shrimp, Penaeus ...
[25]
Effect of eyestalk ablation on ovarian maturation in the tiger shrimp ...
Aug 10, 2025 · Within 3-6 days after eyestalk ablation, 70% of the females matured in the cage and spawned successfully in situ, thereby indicating the ...
[26]
Eyestalk - an overview | ScienceDirect Topics
Male reproductive activity in crustacean decapods is controlled by the X-organ–sinus gland complex (Alfaro-Montoya, 2010). Removal of one (unilateral) eyestalk ...
[27]
OVARIAN MATURATION INDUCED BY EYESTALK ABLATION IN ...
This paper describes attempts to induce maturation of ovaries and ova in pink shrimp (Penaeus duorarum) by eyestalk ablation and by control of environmental ...
[28]
History of shrimp farming - Responsible Seafood Advocate
Sep 1, 2011 · Shrimp farming is a young and dynamic business with a history of rapid change in response to technological advances.
[29]
(PDF) History of Shrimp Farming - ResearchGate
only one eyestalk stimulates maturation without causing mortality. While unilateral eyestalk ablation was sufﬁcient to induce spawning of mature shrimp.
[30]
2. Background
Once nutritional requirements of the broodstock were met, eyestalk ablation techniques led to successful all year reproduction of P. vannamei, and it replaced P ...
[31]
Reproductive performance and offspring quality of non-ablated ...
Mar 30, 2019 · Standard hatchery methods worldwide to control the maturation and reproduction of L. vannamei are based on the use of eyestalk ablation ( ...
[32]
The Shrimp Welfare Dilemma: A Scientific Perspective on Eyestalk Ablation
**Summary of Eyestalk Ablation in Shrimp Aquaculture:**
[33]
Effect of eyestalk ablation on the metabolic activities of two penaeid ...
Prawns in the inter‐moult stage were chosen for eyestalk ablation. During ablation, the eyestalk was removed after holding the prawn in pre‐cooled seawater to ...
[34]
What is eyestalk ablation? - Shrimp Welfare Project
Jul 9, 2025 · The Gonad-Inhibiting Hormone controls shrimps' reproductive organs, when they mature, and when spawning happens. · The Moult-Inhibiting Hormone ...
[35]
Get to Know the Eyestalk Ablation Technique on Shrimp | JALA Blog
Eye stalk ablation is one way to increase the production of shrimp fry. This technique is applied to trigger gonad maturation and brood reproduction.
[36]
GSA Survey Explores the Prevalence of Shrimp Eyestalk Ablation ...
May 20, 2024 · GSA conducted a survey in 2023 about eyestalk ablation among major shrimp producers associations producing two species of shrimp.
[37]
Effect of unilateral and bilateral eyestalk ablation in Litopenaeus ...
Oct 1, 2008 · Males and females were affected differently by eyestalk ablation in terms of concentrations of glucose, triglycerides, and protein in hemolymph.
[38]
The Shrimp Welfare Dilemma: A Scientific Perspective on Eyestalk ...
Aug 18, 2025 · The practice results in a bodily/physical alteration, potential 'pain' as well as physiological impacts on neuro, endocrine, and immune systems ...Missing: effects | Show results with:effects
[39]
[PDF] Effect of unilateral and bilateral eyestalk ablation ... - Archimer - Ifremer
effect of unilateral and bilateral eyestalk ablation ... submitted to unilateral eyestalk ablation in association with a diet supplemented ... crustacean.
[40]
Effects of bilateral and unilateral eyestalk ablation on vitellogenin ...
These results indicate that in immature female prawns, unilateral ablation does not induce VTG gene expression, whereas bilateral ablation induces rapid VTG ...Missing: reproduction | Show results with:reproduction
[41]
Effects of Unilateral versus Bilateral Eyestalk Ablation on Moulting ...
In the present study the moult cycle duration and growth increment were determined in shrimp after unilateral and bilateral eyestalk ablation. Attempts were ...
[42]
Effect of Eyestalk Ablation on Maturation, Larval Performance, and ...
Aug 6, 2025 · We found that females with unilaterally ablated eyestalks showed a better reproductive performance but higher mortality than non-ablated females ...
[43]
[PDF] effects of eyestalk ablation on growth and molt· of freshwater prawn ...
In the group with bilateral eyestalk ablation, the m0lt frequency and growth were greatly accelerated. The group with unilateral eyestalk ablation grew less and.
[44]
None
### Summary of Effects of Eyestalk Ablation on Ovarian Maturation in Macrobrachium lanchesteri
[45]
[PDF] Comparison of ovarian maturation and spawning after unilateral ...
The present study compares the efficiency of ovarian maturation and spawning success between wild-caught and pond-reared. Penaeus monodon females after ...<|control11|><|separator|>
[46]
[PDF] Off-season Maturation and Spawning of the Pacific White Shrimp ...
In the second eyestalk-ablated group (Group 5), a total of 18 females (%90 female spawning rate) spawned generating the highest mean fecundity of 125,015 eggs ...
[47]
Does unilateral eyestalk ablation affect the quality of the larvae of ...
Jun 12, 2018 · This study confirms earlier findings that females with unilaterally ablated eyestalks increase spawn frequency and production of eggs and ...
[48]
Comparison of Reproductive Performance of Domesticated ...
Jul 13, 2020 · In our study, data were collected during the second month after a female had experienced unilateral eyestalk ablation, so production of nauplii ...
[49]
Evaluation of some hemato-immunological parameters in female ...
Mortality was significantly higher in unilaterally (35%) and bilaterally (68%) ablated shrimp than in untreated shrimp (2%), probably caused by impairment of ...
[50]
A feasible path towards year-round production - ScienceDirect.com
The major disadvantages of eyestalk ablation are the high mortality rates recorded for females, during or after surgery, and the irreversible damage caused on ...<|separator|>
[51]
[PDF] The Effect of Eyestalk Ablation on Several Immunologic Variables in ...
The results show that the ablated shrimps' group, after 6 hours, experienced a significant decrease in THC (3.12. ± 0.75 x 106 cells ml-1), but recovered after ...
[52]
Increased robustness of postlarvae and juveniles from non-ablated ...
Although unilateral eyestalk ablation facilitates the establishment of production schedules and increased nauplii production in commercial shrimp hatcheries ...
[53]
Effect of Eyestalk Ablation on Maturation, Larval Performance, and ...
Eyestalk ablation clearly increases reproductive output in Penaeid shrimp, and it is widely used to obtain larvae in commercial hatcheries.Missing: farming | Show results with:farming
[54]
Broodstock management and seed production of Penaeus monodon ...
Induced maturation has mainly been done through the eyestalk ablation method. Nutritional and environmental parameters were studied to enhance reproductive ...<|separator|>
[55]
100 years domestication of penaeid shrimp and meta-analysis of ...
Jun 27, 2024 · Early experiments demonstrated that eyestalk ablations could stimulate ovarian maturation, but female shrimp typically reabsorbed their ovaries ...
[56]
Induced spawning of the mud crab Scylla paramamosain by bilateral...
Eyestalk ablation refers to the removal of one ocular globe from the crustacean's eyestalk to accelerate ovarian maturation. Other techniques, such as hormonal ...<|separator|>
[57]
Stirling researchers identify viable ablation alternatives for shrimp ...
Jan 28, 2019 · Eyestalk ablation– the surgical removal or cutting of a broodstock shrimp's eyestalk – has been commonplace in hatcheries worldwide for the past ...
[58]
Global Shrimp Forum: Shrimp feed demand rises as industry looks ...
Sep 26, 2025 · Estimated farmed shrimp production grew from 4.8 million tons in 2018 to 5.88 million tons in 2024, an increase that has also driven higher ...
[59]
MUTILATIONS | Crustacean Compassion
Eyestalk Ablation of Prawns and Shrimp · Numerous scientists have found that the procedure appears to cause stress, trauma, and pain, with Australian vet Anthony ...Missing: controversies | Show results with:controversies
[60]
Putative Nociceptive Responses in a Decapod Crustacean
Oct 22, 2024 · Nociceptors detect damaging stimuli and send signals to the central nervous system (CNS) about potential injury, which can give rise to pain.
[61]
Minimizing the effects of stress during eyestalk ablation of ...
To induce ovarian maturation and spawning of L. vannamei, unilateral eyestalk ablation is performed on female broodstock (Zaib-Un-Nisa and Ahmed, 2001).<|separator|>
[62]
Pain and stress in crustaceans? - ScienceDirect.com
We consider evidence that crustaceans might experience pain and stress in ways that are analogous to those of vertebrates.
[63]
[PDF] Sentience in decapod crustaceans: A general framework and review ...
May 27, 2022 · This review introduces a framework for evaluating evidence of animal sentience, focussing on pain as an aspect of sentience with particular ...
[64]
Reasons to Be Skeptical about Sentience and Pain in Fishes and ...
Oct 4, 2023 · Applying Mertonian skepticism toward claims for sentience and pain in fishes and aquatic invertebrates is scientifically sound and prudent.<|separator|>
[65]
[PDF] USE OF NON-ABLATED SHRIMP (Litopenaeus vannamei) IN ...
Over the past 4 decades commercial shrimp hatcheries have been using unilateral eyestalk ablation as a method to induce rapid maturation and spawning, hence it ...
[66]
The Shrimp Welfare Dilemma: A Perspective on Eyestalk Ablation
Sep 25, 2025 · Compromised Welfare: Beyond the immediate pain, ablation negatively impacts the overall well-being of the shrimp. Studies indicate that ablated ...
[67]
Eyestalk ablation - Wikipedia
Eyestalk ablation is the removal of one (unilateral) or both (bilateral) eyestalks from a crustacean. It is routinely practiced on female shrimps.
[68]
Research shows shrimp aquaculture industry can move past ...
New research has shown the shrimp aquaculture industry can successfully transition away from eyestalk ablation without adverse affects.
[69]
Identification of structural variation related to spawn capability of ...
Oct 7, 2024 · The goal of shrimp selective breeding is typically to improve economically-relevant traits. ... The female shrimps underwent unilateral eyestalk ...
[70]
Silencing of Gonad-Inhibiting Hormone Transcripts in Litopenaeus ...
This is the first report of the use of RNAi technology to develop an alternative method to eyestalk ablation in captive L. vannamei shrimps.
[71]
Captive maturation studies in Penaeus monodon by GIH silencing ...
Nov 1, 2015 · ... gonad-inhibiting hormone (GIH) secreted from the eyestalk. However, unilateral eyestalk ablation creates physiological complications in the ...
[72]
Silencing of gonad-inhibiting hormone gene expression in Penaeus ...
Aug 10, 2013 · In penaeid shrimp, RNA interference technology was successfully applied to inhibit viral replication as well as to silence shrimp endogenous ...
[73]
Assessment of the Effects of Double-Stranded RNAs Corresponding ...
Mar 2, 2021 · ... gonad-inhibiting hormone, GIH). There now exist many ... eyestalk ablation, but with less stimulatory effect than eyestalk ablation.
[74]
Identification of genes regulated by 20-Hydroxyecdysone in ...
Jan 5, 2024 · 20-hydroxyecdysone (20E) controls various physiological behaviors in crustaceans, among which is the initiation of molt. Previous studies have ...Missing: efficacy | Show results with:efficacy
[75]
Reproductive behavioural differences between wild-caught and ...
Jul 15, 2013 · Unlike the wild shrimps, ovaries of domesticated shrimps do not mature normally without eyestalk ablation. Moreover, domesticated shrimps ...
[76]
Global Seafood Alliance Will Require All BAP-certified Shrimp ...
Aug 27, 2025 · GSA will require all BAP-certified shrimp production facilities to end eyestalk ablation by the end of 2030.
[77]
GSA banning eyestalk ablation in BAP-certified shrimp by end of 2030
The Global Seafood Alliance announced its Best Aquaculture Practices certification standard will no longer allow eyestalk ablation by the end of 2030.
[78]
GSA to ban eyestalk ablation in BAP-certified shrimp by 2030
Aug 28, 2025 · In 2024, the organization conducted a global survey of shrimp producers to evaluate the industry's readiness to move away from ablation and ...
[79]
Ecuadorian shrimp farms ditch eyestalk ablation - LinkedIn
Aug 31, 2025 · Most Ecuadorian broodstock facilities, such as Biogemar, Aquagen, Omarsa, and others, have completely eliminated eyestalk ablation in female ...Missing: crustacean Southeast Asia
[80]
International seafood certification standards on eyestalk ablation
The standard requires that from no later than April 2024, larvae may only be sourced from shrimp females which have not suffered eye ablation (or any other ...
[81]
Co-op is the latest UK supermarket to end prawn eyestalk ablation
Co-op has been working to eliminate eyestalk ablation from its supply chain since 2022, when 72 percent of their female prawns were non-ablated. By 2023, that ...<|control11|><|separator|>
[82]
Biosecurity along the shrimp value chain - INVE Aquaculture
Oct 26, 2023 · In both trials, the offspring from non-ablated females had greater rates of survival, significantly so when challenged with an isolate of ...Missing: studies | Show results with:studies