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Review

Evaluating Alternative and Sustainable Food Resources: A Review of the Nutritional Composition of Myctophid Fishes

by
Bowen Zhang
1,2,*,
Heidi Pethybridge
2,
Patti Virtue
1,2,
Peter D. Nichols
1,2,
Kerrie Swadling
1,
Alan Williams
2 and
Kim Lee-Chang
2
1
Institute for Marine and Antarctic Studies, University of Tasmania, Battery Point, TAS 7004, Australia
2
CSIRO Environment, Battery Point, TAS 7004, Australia
*
Author to whom correspondence should be addressed.
Sustainability 2023, 15(15), 12039; https://doi.org/10.3390/su151512039
Submission received: 16 May 2023 / Revised: 28 July 2023 / Accepted: 4 August 2023 / Published: 6 August 2023
(This article belongs to the Special Issue Marine Biotechnology for Sustainability of Ecologically Significant Resources)
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Abstract

:
Additional and alternative sustainable food resources are needed as the global human population increases. Marine fishes have long provided essential nutrients, such as omega-3 long-chain (≥C20) polyunsaturated fatty acids (n-3 LC-PUFA), protein, and vitamins to meet human dietary requirements and feed for agricultural production. Many current commercial fish stocks are depleted or fully exploited, but oceanic mesopelagic fishes, particularly the myctophids (lanternfishes), represent a potentially very large and unfished resource. This review analysed the literature on nutritional and biochemical compositions of myctophids as a first step towards understanding the health benefits and risks of consuming them. We found that myctophids have high levels of protein (11–23% wet weight, WW) and variable lipid content (0.5–26% WW). In most species, desirable triacylglycerols or phospholipids dominated over less-desirable wax esters, and most have abundant amounts of health-promoting n-3 LC-PUFA, such as DHA and EPA. Myctophids have low levels of heavy metals and persistent organic pollutants. Most nutritional information is available for species from the Pacific and Southern Oceans and for the genera Benthosema, Electrona, and Diaphus. Myctophids generally possess favourable nutritional profiles, but major gaps in knowledge regarding their stock assessment, ecology and the economic viability for their harvest are barriers to developing sustainable fisheries.

1. Introduction

As the global human population increases, the demand for food and feed resources is escalating proportionally. Marine fishes are food with exceptional nutritional value that has been recognised for their positive impact on human health [1,2]. It has been reported that fish consumption represents about 20% of human intake of animal protein worldwide. The high-quality proteins in fish contain many essential amino acids that are vital for human growth and maintenance [3]. Fish constitutes a major dietary source of polyunsaturated fatty acids (PUFAs) [4], particularly the omega-3 (n-3) long-chain (≥C20) polyunsaturated fatty acids (LC-PUFA, also termed LC omega-3 oils), including eicosapentaenoic acid (EPA, 20:5n-3) and docosahexaenoic acid (DHA, 22:6n-3). These essential fatty acids are crucial to human brain development and contribute to the maintenance of cardiovascular health [5,6], and can provide protection against coronary heart disease and ischemic stroke. Fish is also considered a valuable dietary source of micronutrients, such as vitamin D, calcium, selenium, iodine, and zinc [7], which are essential for maintaining physiological function [8].
Worldwide, many coastal demersal and pelagic fisheries are at maximum harvest capacity and, in some instances facing the threat of collapse [9,10]. The decline in overall biomass can be attributed to a variety of factors, including overfishing, habitat destruction, pollution, and, more recently, climate change [11,12]. Declining fish stock and biomass can have cascading effects through the food chain and alter the structure and functioning of entire ecosystems [12,13]. Such ecological changes affect not only the amount of food available for human consumption, but also the nutritional quality. Growing concerns for current coastal and terrestrial food systems mean that new sources of nutrition are needed.
The mesopelagic zone (200–1000 m) contains a large diversity and abundance of small, migratory organisms that serve a key role in regulating the carbon and nutrient cycles globally [14,15]. Myctophid fishes are among the most abundant and widely distributed taxa, as well as one of the most diverse families of the teleost fishes [16], with over 250 species in 33 genera [17] (one species is illustrated in Figure 1). While there are order-of-magnitude disparities in the estimates of the worldwide biomass of mesopelagic fishes, the resource is prospectively very large at 1 to ~16 Gt (109 tonnes) [17,18,19,20,21,22] and made up mostly of myctophids. These data suggest that myctophids could potentially contribute to meeting the increasing global demand for nutrients derived from marine sources. Myctophids are also thought to be a promising future source of health-benefiting LC omega-3 oils, protein, and essential minerals and vitamins [23]. As a mid-trophic and oceanic resource, they are reported to have low levels of toxins and heavy metals, including mercury, lead, arsenic, and cadmium [23]. There are, however, a number of reports which indicate that some myctophids can have undesirable features (e.g., high wax esters; see Section 2.1) and that there are challenges associated with processing myctophids on a large scale which may hinder their use as a food source [24,25].
The primary aims of this review were to (i) examine the nutritional composition of myctophid fishes caught in different oceans to assess their suitability for human consumption; and (ii) summarise aspects of myctophid ecology, including knowledge gaps relevant to the future exploitation of myctophid resources. For each nutritional component, we summarise the available data in the most widely recognised unit of analysis (typically expressed in wet mass as a concentration or relative percentage) and for whole specimens of species.

2. Nutritional Profiles

2.1. Lipids & Fatty Acids

Lipids are the densest form of stored energy and provide at least two-thirds more energy per gram than proteins or carbohydrates [26]. The need for fish oil in recent years has grown mostly due to growth in the aquafeed (aquaculture) and nutraceutical industries [27,28]. The price of fish oil has increased from around 750 USD/ton to 3000 USD/ton over the last 15 years [29,30]. Between 2009 and 2013, there was a considerable increase in the real prices of fish oil, which subsequently declined until 2017, although have since increased again [30]. The fish oil price has increased by 45.1% in the past decade, and fish oil production will grow by 0.8% p.a. by 2030 [30]. Global prices for fishmeal have also increased, athough they are highly variable and dependent on the amount of oil (lipid) and its composition [30]. The most desirable characteristics of these fish oils are high amounts of triacylglycerols (TAG) and n-3 LC-PUFA, particularly DHA and EPA, which are essential dietary components for human and animal health.
The lipid class composition of fish oil is also an important feature, as they have distinct effects on human and animal health. Certain types of fats (e.g., TAG-based saturated and trans fats) can increase the risk of heart disease and other health problems [31,32]. Wax ester (WE) is a class of lipids that are considered less desirable for human consumption due to their propensity to cause keriorrhea, a digestive disorder characterised by diarrhoea and abdominal discomfort, and limited digestibility [16,33]. However, WE have potential applications in various industries, including cosmetics (such as occurs for jojoba oil [23]), food (such as the coating material [34]), and biofuel industries [35].
Our assessment of the literature showed that the lipid content in whole individuals of myctophid species varies widely from 0.5 to 26% wet weight (WW), with those characterised by high lipid content (>10% WW) and dominated by TAG considered most desirable for food nutrition (Table 1). Such high variations between species have been reported in regional studies (e.g., [36] showed a range of 4.4–26.1%). Some species have high lipid content (>10%), which is higher than many other commercially valuable fish, including mackerel icefish [24] and capelin, sand lance, and squid in the North Pacific [37]. To date, few studies have examined the influence of geographical location on the lipid content and composition of myctophid species [24]. In a regional comparison study, significant differences in the lipid composition of myctophids were reported between those sampled in the Indian and Pacific sectors of the Southern Ocean [24]. In a study characterising the lipid of 20 myctophid species in both the subarctic and tropical Pacific Ocean, it was found that those sampled in tropical waters had much less lipid and were typically not dominated by WE or TAG [38]. Our assessment of the available literature also suggests that myctophids harvested in polar regions mostly have intermediate to high lipid content (>5% WW), with only one species, Gymnoscopelus fraseri from Macquarie Ridge, reported to have low lipid content (3.6% average WW) [24]. In contrast, there is more variability in lipid content reported between species of myctophids from temperate and tropical ecosystems; this seems to be attributed to phylogeny in addition to geographical location and perhaps time of capture. For one of the most studied genera of myctophids, Diaphus, we found that lipid content varies considerably with examples of both high and low lipid content in almost all ocean bodies assessed, but the lipid composition is consistently dominated by TAG. We found that no studies have explored long-term (decadal) trends in the lipid content of any myctophid species. Some studies have explored short-term temporal variability in the lipid compositions of myctophids noting seasonal [39] and interannual differences [38].
Myctophids have variable lipid class compositions and typically are either TAG or WE dominant (Table 1), although many species have substantial amounts of phospholipids (PL) [38,40]. Except for several species (e.g., Electrona antarctica, Figure 1), which have a high WE content [41], TAG is the dominant lipid class in most of the Southern Ocean myctophid species studied to date [24,33,42,43,44]. In temperate and tropical myctophids, lipid class composition seems to differ among tax; fewer species were dominated by either WE or TAG, but rather lipid class profiles consisted of a combination of TAG and PL [36,38,40]. A spatial comparison study looked at the effect of vertical distribution and migration and suggested that myctophids that are WE-dominated do not undergo large vertical migrations [38]. Species that are dominant in WE (Table 1) may be a problem for both direct human consumption and for use in aquafeeds, as most humans and many fish species have limited capacity to utilise them [16].
Our analysis of literature reporting fatty acid composition for whole myctophids found that they have higher levels of DHA than EPA and that there are large variations between species and study regions (Table 2). Across all the study regions, levels of EPA ranged from 0 to 1042 mg/100 g (0 to 7.7% of total fatty acids) WW, while DHA ranged from 298 to 2016 mg/100 g (2.1 to 25.1%) WW. Genera with the highest relative amounts of DHA include a Diaphus species in the subantarctic Pacific Ocean [38] and Gymnoscopelus fraseri in the Pacific section of the Southern Ocean [24]. Species highest in relative levels of EPA included Lampanyctus australis [40] from the South Pacific Ocean. Most studies show that myctophids are dominated by monosaturated fatty acids (MUFA), such as 18:1 and 22:1. There is evidence to suggest that the fatty acid composition of myctophids is strongly aligned with the lipid class [24,33,42]. Higher relative levels of EPA were found in the WE–rich species, while DHA was higher in TAG–rich species [33,38,41]. Palmitic acid (16:0) and DHA also seem to be more abundant in myctophid species with high levels of PL [45].
Table 1. Overview of the total lipid content and TAG & WE dominance in myctophid taxonomic groups (up to genus level) in relation to oceanic regions. Lipid content is expressed as a percentage of wet weight (% WW), while TAG & WE are expressed as a percentage of total lipid content. Dominance classifications are based on mean reported values. References for taxa with moderate levels of total lipid content (5–10% WW) reported are not included.
Table 1. Overview of the total lipid content and TAG & WE dominance in myctophid taxonomic groups (up to genus level) in relation to oceanic regions. Lipid content is expressed as a percentage of wet weight (% WW), while TAG & WE are expressed as a percentage of total lipid content. Dominance classifications are based on mean reported values. References for taxa with moderate levels of total lipid content (5–10% WW) reported are not included.
LocationGeneraCatch YearRefs.
High lipid (>10% WW)Indian OceanDiaphus2009[46]
Diaphus2011[47] ^
Pacific OceanDiaphus2012[39]
Ceratoscopelus, Lampadena, Lampanyctus,
Notoscopelus, Protomyctophum, Stenobrachius,
Symbolophorus		1994[38]
Ceratoscopelus, Notoscopelus, Symbolophorus,
Diaphus, Myctophum, Lampanyctus,
Protomyctophum, Stenobrachius		1995[36]
Lampanyctus, Triphoturus, Symbolophorus1975–1979[48]
Diaphus, Protomyctophum, Symbolophorus2005[40]
Atlantic OceanBenthosema2015–2018[49,50]
SubarcticDiaphus, Lampanyctus, Notoscopelus, Stenobrachius, Symbolophorus1992–1994[38]
Southern OceanElectrona, Gymnoscopelus, Krefftichthys,
Protomyctophum		1995[42]
Metelectrona1987[51]
Electrona, Gymnoscopelus1999, 2008[24,43]
Electrona2009–2012[44]
Low lipid
(<5% WW)		Indian OceanMyctophum2012[47] ^
Diaphus, Benthosema, Myctophum2013[52]
Pacific OceanBenthosema, Diaphus2012[39]
Diaphus1994[38]
Benthosema, Ceratoscopelus, Diaphus,
Hygophum, Lampadena, Myctophum		1993[38]
Lampanyctus1995[36]
Bolinichthys, Diaphus, Gonichthys, Hygophum, Myctophum, Notoscopelus, Protomyctophum, Taaningichthys, Tarletonbeania, Lampanyctus, Ceratoscopelus1975–1979[48]
Diaphus, Electrona, Hygophum,
Lampanyctus, Lampanyctodes, Lampichthys,
Metelectrona, Nannobrachium		2005[40]
Atlantic OceanLampanyctus, Hygophum1968[53]
Southern OceanGymnoscopelus1999[24]
TAG dominant (>60%)Pacific OceanCeratoscopelus, Symbolophorus1993[38]
Bolinichthys, Lampanyctus, Symbolophorus1975–1979[48]
Diaphus, Lampadena, Lampanyctus,
Lampichthys, Notoscopelus, Protomyctophum		1994[38]
Diaphus, Lampanyctus, Notoscopelus,
Symbolophorus		2005[40]
Benthosema, Diaphus2012[39,47] ^
Indian OceanMyctophum, Diaphus2011[47] ^
SubarcticDiaphus, Lampanyctus, Symbolophorus1992[38]
Southern OceanElectrona, Gymnoscopelus1999[24,33]
WE dominant (>40%)Pacific OceanLampanyctus, Stenobrachius1994[38]
Lampanyctus, Triphoturus1975–1979[48]
Nannobrachium2005[40]
Atlantic OceanBenthosema2015–2016[50]
SubarcticLampanyctus, Stenobrachius1992[38]
Southern OceanElectrona, Gymnoscopelus, Krefftichthys1999[24,33,42]
^ myctophid fish in this study were filleted, deskinned and homogenised for lipid analysis.
Table 2. Reported ranges of mean values of EPA and DHA (as % of total fatty acids, wet weight) in the whole body of myctophid species studied in different regions.
Table 2. Reported ranges of mean values of EPA and DHA (as % of total fatty acids, wet weight) in the whole body of myctophid species studied in different regions.
OceanRegion# Species AssessedEPADHAReference
IndianArabian Sea13.89.3[46]
Arabian Sea44.1–7.07.6–20.2[52]
Arabian Sea34.3–5.89.8–15.9[47] ^
PacificPapua New Guinea90–1.53.8–10.3[38]
Subantarctic170–2.210.1–23.9[38]
Coast of Japan30.9–1.56.9–18.5[39]
Tasman Sea123.0–7.27.4–19.8[40]
AtlanticNorwegian fjords16.210.4[49]
North-eastern41.0–4.82.1–17.34[54]
Southern OceanKerguelen Plateau60.7–1.15.5–12.2[53]
Macquarie Island21.2–1.418.9–20.5[53]
Kerguelen50.3–4.63.9–7.4[42]
Antarctic Peninsula11.7–6.92.9–8.8[41]
Heard Island154.1–7.76.3–17.0[55]
^ myctophid fish in this study were filleted, deskinned and homogenised for lipid analysis.

2.2. Proteins & Amino Acids

Marine fish have long been recognised as a good source of high-quality protein that contains all the essential amino acids required for optimal health benefits [56,57,58,59]. Protein content is also an important factor in determining economic value [60,61]. Protein derivatives from fish, including mince, surimi, protein isolate, hydrolysate, and powder, have been used in the food industry [23]. From the available literature (Table 3), we found that the total protein content of myctophids varies from 11% to 23% (WW) [23,25,62,63,64,65,66,67], which has been shown to be similar or higher than other commercial fish and shellfish [68]. Differences between myctophid species have been noted [23,63,64,66], especially in the two highly abundant genera, Diaphus and Lampanyctus. For instance, Diaphus hudsoni (off the south-west coast of India, [66]) and Diaphus watasi (Arabian Sea, [65]) exhibit protein contents of 14% and 21% (WW), respectively, while Diaphus luetkeni and Diaphus effulgens both have protein contents of 16.5% (WW) (Arabian Sea, [69]). The majority of protein research conducted on myctophids (Table 3) has been focused on the Indian Ocean, such as the Gulf of Oman [63] and Arabian Sea [23,52,65], and the western Pacific (southwestern Taiwan) [67]. However, a large knowledge gap exists for myctophid species inhabiting other oceanic regions, where limited research has been carried out.
The amino acid composition of 13 myctophid species, expressed as a percentage of protein (g/100 g of total protein), has been reported, with total amino acid content ranging from 40% to 52% WW (Figure 2) [25,62,63,64,65]. No significant differences were observed between the amino acid content of migratory and non-migratory myctophids from the subarctic and tropical Pacific Ocean [64]. Essential amino acids, including valine, methionine, leucine, isoleucine, tyrosine, phenylalanine, histidine, lysine, threonine, and tryptophan, exhibited the highest concentrations in Diaphus effulgens (52%), followed by Diaphus watasei (47%), Benthosema fibulatum (44%), Myctophum obtusirostre (43%), Diaphus hudsoni (42%), and Diaphus luetkeni (39%) [25,62,63,64,65]. Leucine, lysine and arginine, which play a vital role in calcium absorption and metabolism [70,71] in humans, are the most dominant amino acids in myctophids (Figure 2), while histidine is also high in Diaphus effulgens [66].
Table 3. Overview of protein content (percentage of wet weight; WW) of myctophids from the Indian, Pacific and Atlantic Oceans.
Table 3. Overview of protein content (percentage of wet weight; WW) of myctophids from the Indian, Pacific and Atlantic Oceans.
OceanGenusProtein (% WW)Catch YearReference
IndianBenthosema16.1–18.6 [23,52]
Ceratoscopelus11.51979[63]
Diaphus13.3–21.41979, 2009, 2012[46,52,63,65,66]
Lampadena11.51979[63]
Lampanyctus12.1–13.41979[63]
Myctophum19.3–22.3 [52,65]
Notoscopelus13.51979[63]
Stenobrachius12.5–12.81979[63]
Symbolophorus12.31979[63]
Western PacificBenthosema14.4–15.0 [67]
Northern AtlanticBenthosema41.12015–2016[50]

2.3. Minerals and Vitamins

Fish are considered a vital source of dietary essential minerals and trace elements [72,73,74,75], which are more readily absorbed and utilised by humans in comparison to alternative food sources. This includes calcium, which is fundamental to skeletal growth and various physiological functions [50], as well as magnesium, a key nutrient required for metabolic processes [54]. Moreover, iron is an essential mineral that is critical for the production of red blood cells and oxygen transport in fish [1]. Myctophids sampled in the northern Indian Ocean have higher levels of calcium (>900 mg/100 g), phosphorus (500 mg/100 g), potassium (>300 mg/100 g), and iodine (>100 µg/100 g), compared with small pelagic fish (such as Fringescale sardinella) and benthic fish species (such as Bombay duck, Harpadon nehereus) [76]. Among known edible species of myctophids, Diaphus watasei has been identified as having a relatively high mineral content, specifically in terms of potassium and calcium, with concentrations of 35.3 and 47.3 mg/100 g, respectively [69]. Another abundant species, Benthosema glaciale, in the Northern Atlantic also has been reported to have good levels of favourable macro minerals and trace elements [50].
Table 4 presents an overview of the macro minerals and trace elements found in myctophid species, including Benthosema glaciale [50,76] and Benthosema fibulatum [76]. The macro minerals analysed included calcium, phosphorus, magnesium, sodium, and potassium, while the trace elements included iron, manganese, zinc, copper, selenium, nickel, lead, and iodine. The results show that Benthosema glaciale has higher levels of calcium, phosphorus, and potassium, while Benthosema fibulatum has higher levels of magnesium, copper, and iodine. The recommended intake (RI) values suggest that these myctophid species can be a valuable source of these essential minerals and trace elements for human consumption, especially for meeting dietary requirements for calcium, phosphorus, and iodine [77]. The levels of trace elements found in both species fall within the acceptable ranges recommended by health organisations, making them safe for human consumption.
Fish is also considered an excellent source of soluble vitamins, including vitamins A, D, and E, in addition to vitamin B12 which are important for energy production, functioning of the nervous systems, and metabolic processes [78,79]. Myctophids can provide a rich source of soluble vitamins, particularly vitamins A and E [47,80]. Past research suggests that myctophids have higher levels of vitamin A1 (>100 µg/100 g raw, edible part) and vitamin B12 (6.2 µg/100 g raw, edible part on average), while vitamin A2 and D were found to be significantly lower in demersal fish (<0.5 µg/100 g raw, edible part) [76]. A study conducted on myctophids in the Norwegian fjords also revealed very high levels of vitamin A, particularly in the head and viscera [49]. Despite their potential as a nutrient-rich food source, the vitamin composition of myctophids remains poorly documented, and more comprehensive research is required to assess their potential, both for direct human consumption and as feed ingredients in aquaculture.
Table 4. Summary of macro minerals: calcium (Ca), phosphorus (P), magnesium (Mg), sodium (Na), potassium (K), and trace elements: iron (Fe), manganese (Mn), zinc (Zn), copper (Cu), selenium (Se), nickel (Ni), lead (Pb), iodine (I) in myctophid species (Benthosema glaciale and Benthosema fibulatum). The recommended intake (RI) per day for these minerals and trace elements is also provided [77].
Table 4. Summary of macro minerals: calcium (Ca), phosphorus (P), magnesium (Mg), sodium (Na), potassium (K), and trace elements: iron (Fe), manganese (Mn), zinc (Zn), copper (Cu), selenium (Se), nickel (Ni), lead (Pb), iodine (I) in myctophid species (Benthosema glaciale and Benthosema fibulatum). The recommended intake (RI) per day for these minerals and trace elements is also provided [77].
Benthosema glacialeBenthosema fibulatumRI (mg)
Catch year2015, 2016201820182018
Ocean LocationAtlantic Indian AtlanticIndian
Reference[50][76][49][76][77]
Macro minerals (mg/g DW)Ca12.1 5.09.4540–900
P8.3 3.85.8420–700
Mg1.2 0.70.680–350
Na5.4 3.92.3
K7.5 2.63.02900–3510
Trace elements (mg/kg DW)Fe51.6 10.825.05–15
Mn2.8
Zn36.0 8.015.05–12
Cu3.0 0.3–1.0
Se1.70.060.60.10.015–0.06
Ni0.9
Pb0.4
I 0.04 0.20.05–0.15

2.4. Contaminants

Knowledge about contaminants in any food source is important for a proper assessment of potential health risks associated with human consumption, regulatory compliance, and consumer awareness. Food safety regulations, as dictated by the European Union or the US food safety regulations, have been set particularly tight according to recommended intake levels. Reporting of contaminant concentrations in myctophids is presently limited, but is available for toxic metals (e.g., mercury) [81,82,83,84,85] (Table 5) and plastic-associated chemicals (e.g., bisphenol A (BPA), alkylphenol ethoxylates (APEs), pesticides, polychlorinated biphenyls (PCBs), and polybrominated diphenyl ethers (PBDEs)) [86,87,88,89,90,91,92,93] (Table 6).
From our assessment of the literature, ten regional studies have reported total mercury (THg) concentrations in myctophid species from the Atlantic, Southern and Pacific Oceans. This research has provided evidence that myctophids contain very low levels of Hg, with mean values consistently lower than 0.5 μg/g DW (Table 5), which is below the tolerable daily intake (TDI) of 0.05 mg per day [94]. THg ranges in myctophids are similar across the globe ranging between 0.03 and 0.45 μg/g DW in the North Atlantic Ocean [95,96,97,98,99], 0.03 and 0.42 μg/g DW in the Southern Ocean [82,84,100], and between 0.04 to 0.32 μg/g in the Southwest Pacific Ocean [40]. Among the genera studied, the highest Hg concentrations were observed in species from the genera Benthosema, Ceratoscopelus, and Gymnoscopelus from the Atlantic Ocean and the Southern Ocean. A study in the North Atlantic Ocean showed that Hg levels in an abundant myctophid species, Benthosema glaciale, had not changed between 1936 to 1993, but that there was significant regional differences [98]. Other regional studies have shown evidence that inter-specific variations in THg levels of myctophid are linked to size [83] and vertical distribution [101], with higher levels reported in larger and deeper dwelling species. We found only one study that has reported the proportion of THg that consists of the toxic methylmercury form in myctophids from the Mediterranean Sea and indicated that it ranges from ~70–82% [102].
Table 5. Summary of total mercury (THg) concentration range (ug/g dry weight; DW) in myctophids of various genera from different oceanic regions. * Values were converted from wet to dry weight, assuming a water content of 75%.
Table 5. Summary of total mercury (THg) concentration range (ug/g dry weight; DW) in myctophids of various genera from different oceanic regions. * Values were converted from wet to dry weight, assuming a water content of 75%.
GenusTHgLocationCatch YearReference
Benthosema0.11–0.45NW Atlantic Ocean1936–1993[98]
Bolinichthys0.02–0.2N Pacific Ocean,2007–2011[95]
0.16NW Atlantic Ocean1971[95,96]
Ceratoscopelus0.21–0.42N Atlantic Ocean1971, 1978[96,97]
Diaphus0.10–0.11NW Atlantic Ocean1971–1974[95,96]
Electrona0.05–0.27Southern Indian Ocean,
Southern Ocean		1997–1999, 2007–2013, 1995[82,84,97,100]
0.08–0.20 *SW Pacific Ocean2005–2006[40]
Gymnoscopelus0.06–0.424Southern Ocean2015–2016, 2007–2011, 1997–1998[82,84,100]
Hygophum0.25–0.30NW Atlantic Ocean1973–1974[95,96]
0.08–0.16 *SW Pacific Ocean2005–2006[40]
Krefftichthys0.03–0.05Southern Ocean2007–2008[84]
Lampanyctus0.16–0.34NW Atlantic Ocean1974[96]
0.12–0.28 *SW Pacific Ocean2005–2006[40]
Lampichthys0.08–0.20 *SW Pacific Ocean2005–2006[40]
Lobianchia0.20–0.24NW Atlantic Ocean1973–1974[95,96]
Myctophum0.08–0.32N Atlantic Ocean1994, 2001–2010[97,103]
Nannobranchium0.28–0.32 *SW Pacific Ocean2005–2006[40]
Notoscopelus0.03–0.24NW, NE Atlantic Ocean1974, 2001–2003[96,99]
0.08–0.12 *SW Pacific Ocean2005–2006[40]
Protomyctophum0.06–0.10S Indian Ocean,
Southern Ocean		1997–1999, 2007–2009[84,100]
Symbolphorus0.04–0.24 *SW Pacific Ocean2005–2006[40]
We reviewed five scientific studies that have examined concentrations of plastic-associated chemicals in myctophids (Table 6). These studies show that for most congeners, concentrations range from non-detectable amounts to levels that are above guidelines and tolerable daily intake levels. Between different species, the reported ranges of the concentrations of bisphenol A (BPA) and alkylphenols, alkylphenol ethoxylates (APEs) are comparable, while there seems to be more between species variability for alkylphenols. Total polychlorinated biphenyls (PCBs) concentrations range between 0.02–5.97 ng/g WW, with the highest reported levels in Myctophum species from the western North Pacific [90]. Reported levels of total polybrominated diphenyl ethers (PBDEs) are highly variable in the North Pacific Gyre, while notably lower in other regional studies.
Table 6. Comparison of plastic concentrations (ng/g wet weight; WW), including bisphenol A (BPA), alkylphenols, alkylphenol ethoxylates (APEs), polychlorinated biphenyls (PCBs), and polybrominated diphenyl ethers (PBDEs), in myctophids from different oceans and the corresponding tolerable daily intake (TDI) (nd is non-detectable).
Table 6. Comparison of plastic concentrations (ng/g wet weight; WW), including bisphenol A (BPA), alkylphenols, alkylphenol ethoxylates (APEs), polychlorinated biphenyls (PCBs), and polybrominated diphenyl ethers (PBDEs), in myctophids from different oceans and the corresponding tolerable daily intake (TDI) (nd is non-detectable).
LocationNorth Pacific OceanWestern North Atlantic OceanNorth Pacific GyreSouthern OceanTDI
SpeciesMyctophum nitidulumTaaningichthys crenularisNot specifiedGymnoscopelus nicholsi
Catch Year2009200920102003
BPAnd-5.10nd-7.70nd-6.20 4 ng/g [104]
Alkylphenolsnd-47nd-62nd-10.80
APEsnd-11nd-8.70nd-5.40
PCBs0.55–3.400.84–1.300.02–5.97 1–4 ng/g [105]
PBDEs0.01–0.050.05–0.590.002–11.200.09 ± 0.020.15 ng/g [106]
Reference[90][90][87][91]

3. Considerations for Future Exploitation of Myctophids

The above section reviewed literature reporting nutritional information for myctophids from around the globe. While the information suggests that many myctophid species can provide a nutritionally valuable alternative food resource for human consumption or agriculture production, other factors will determine the viability of fisheries to exploit them. Here we briefly summarise some of the knowledge of myctophid ecology that is relevant to developing fisheries for myctophids and other mesopelagic fishes: identifying suitable target species, biodiversity and distribution, biomass estimates, growth and life history, and the ecosystem.

3.1. Species Diversity and Distribution

Our review demonstrates important interspecies differences in the nutritional composition of myctophids, indicating that any potential future exploitation may need to consider relevant factors at the species level. This may be challenging because, with 250 recorded species, the Family Myctophidae is one of the most diverse fish families in the world’s oceans [16]. Several regional surveys and studies have reported on species diversity: in the Indian Ocean, 137 myctophid species in 28 genera have been reported, with the most abundant species from the genera Diaphus and Lampanyctus [107]. In areas of the eastern tropical northern Atlantic Ocean, over 80 species of myctophids have been identified, with the genera Benthosema, Myctophum, Diaphus, Lampanyctus, and Hygophum most prevalent [107]. In parts of the Pacific Ocean, approximately 60 species of myctophids are recorded to occur: Myctophum asperum, Diaphus theta, and Lampanyctus festivus are particularly abundant in the western Pacific; Ceratoscopelus warmingii, Hygophum hygomii, and Diaphus spp. are abundant in the southern region [108,109] and Hygophum hygomii in the northern region [16,110,111]. In the Southern Ocean, abundant species of myctophids include Electrona antarctica (Figure 1), Gymnoscopelus braueri, G. ophisthopetrus, G. nicholsi, Krefftichthys anderssoni, Protomyctophum bolini and P. tensioni [24].
A distinct characteristic of many myctophid species is their diel vertical migration. This phenomenon describes night-time ascent by large fractions of the populations of many migratory species from their daytime residence depths below 200 m (mostly below 400 m) to surface or near-surface depths (0–100 m) to feed [17,112,113]. Patterns of migration vary between species and locations with factors, including life history stage, sex, latitude, hydrography, topography, and season believed to influence them [111,114,115]. While strong currents and eddies may affect the horizontal distributions of mesopelagic fishes at local or regional scales, their distributions generally coincide with those of major water masses [17]. This interplay between environmental factors and inherent biological characteristics shapes both the migratory behaviour and biomass distribution of myctophids [17,112,113] and the suitability of environmental habitats [101,107,108,109,116]. These regional, seasonal and daily patterns of habitat use, which are often species-specific, have important implications for the availability of myctophids to fishing fleets.

3.2. Life History and Growth

The majority of myctophids, in common with numerous other marine fish species, are presumed to be broadcast spawners, reproducing via the release of planktonic eggs and larvae that drift with ocean currents, where fertilisation takes place [117]. Both male and female myctophids are non-guarding pelagic spawners, with females being oviparous [16]. Various myctophid species display sexual dimorphism and size disparities, with some smaller myctophid species exhibiting equal sex ratios and no size differences between the sexes [17,62,118,119]. As myctophid larvae mature, they transition from the epipelagic zone (0–200 m) to deeper waters. These young fish possess the ability to navigate density gradients, including the thermocline and halocline, which typically impede mixing through physical processes [120]. In tropical and temperate waters, reproductive activity can vary seasonally, contributing to the variation observed in body size and individual biomass [121].
Myctophids are generally short-lived compared to many deep-sea and coastal fish species; their lifespans range from less than a year to 8 years, depending on the species and geographical locality [17,118,119,122] (Table 7). Daily growth increments in otoliths have been utilised to confirm the cyclical pattern of vertical migration in various species, with most migratory myctophids exhibiting a standard asymptotic growth curve [123]. There is a correlation between growth rate and vertical migration behaviour, with daily vertically migrating species exhibiting comparatively slower growth rates than non-migratory species [122]. Maximum age varies with genera; for example, species of Lampanyctus have been documented to reach a maximum age of 4.5 years in the Pacific Ocean [118,124], while Lampanyctus species in the Atlantic Ocean can reach 5.5 to 6 years [118,125]. Short-lived species may be suited to commercial harvest, but the substantial influence of the surrounding marine environment, likely temperature, on growth and longevity has direct implications for potential fishery productivity.
Table 7. Overview of maximum age (yr: year; d: day) and geographical distribution of selected myctophids from the available literature.
Table 7. Overview of maximum age (yr: year; d: day) and geographical distribution of selected myctophids from the available literature.
GeneraMax AgeOcean RegionRef.
<2 yrBenthosema; Diaphus; Lepidophanes325 d; 362 d; 439 dAtlantic Ocean[126]
Ceratoscopelus; Stenobrachius416 d; 541 dPacific Ocean[127,128]
Myctophum440 d[129]
Tarletonbeani504 d[130]
Benthosema~1 yrIndian Ocean[131]
Myctophum1 yrAtlantic Ocean[132]
Protomyctophum1.25 yr[133]
2–4 yrCeratoscopelus2 yrAtlantic Ocean[134]
Kreffichthys2 yrSouthern Ocean[125]
Diaphus2.5 yrPacific Ocean[135]
Lampanyctodes3 yr[136]
Electrona3.5 yrSouthern Ocean[137]
4–6 yrBenthosema4.5 yrAtlantic Ocean[138]
Lampanyctus; Triphoturus4.5 yr; 5 yrPacific Ocean[118,124]
Benthosema5 yrAtlantic Ocean[131]
Lampanyctus5.5–6 yr[118,125]
6–8 yrGymnoscopelus6 yrSouthern Ocean[125]
Benthosema, Gymnoscopelus7 yrAtlantic Ocean[139,140]
Benthosema7 yrPacific Ocean[141]
Stenobrachius7.5 yrAtlantic Ocean[118]
Stenobrachius8 yrAtlantic Ocean[142]

3.3. Role in the Ecosystem

The ecological characteristics and trophic interactions of myctophids and the importance of their roles within the mesopelagic ecosystem have been long recognised [143,144], but many key knowledge gaps remain. Myctophids have been identified as key prey items of commercially important species, including tropical tunas [145,146] and Southern Ocean toothfish [147]. In the northeastern Pacific, dolphins have been reported to consume significant amounts of myctophids [148]. In the Southern Ocean, myctophids contribute a large proportion of the diet of squid [149], penguins and other seabirds [150,151,152] and marine mammals [143]. Despite the established importance of myctophids in other ecosystems, there is currently limited information available on their ecological role within the subarctic or within tropical regions.
As critical components of marine food webs, myctophids bridge the gap between primary producers and higher trophic-level predators by consuming mesozooplankton [36,119,152,153]. Myctophids exhibit opportunistic predation on a diverse array of prey, including copepods, ostracods, euphausiids, hyperiid amphipods, chaetognaths, pteropods, fish eggs, and fish larvae [17,131]. There is evidence to suggest that diet changes with size and age; for example, early-stage Electrona antarctica feed on sinking particles [154], copepods and hyperiid amphipods, while the larger fish feed on euphausiids [155]. Despite their ecological importance, the full extent of trophic interactions and dietary preferences of most myctophid species are not yet comprehensively understood. In addition to their contribution to the food web, these mesopelagic fish are integral to biogeochemical cycling in the open ocean [25]. Through diel vertical migration and the generation of sinking faeces, myctophids provide a conduit for the significant export of nutrients, trace elements and organic matter between the surface and deep ocean [156]. The many vital ecological roles of myctophids and their importance in the diets of higher-order and often charismatic predators, together with many knowledge gaps, indicate there will be complex environmental challenges to operating and managing future fisheries according to principles of ecological sustainability.

3.4. Biomass and Fishery Potential

The global biomass of mesopelagic animals, including myctophids, has been estimated using several methods: pelagic trawl surveys [17], acoustic surveys [18,19,157], modelling [20,21,22] and remotely operated vehicles [158,159,160]. The first (in 1980) [17] was based on a compilation of trawl survey data from many ocean regions. It estimated total mesopelagic fish biomass to be approximately 1 Gt; myctophids formed a large fraction of biomass in all regions surveyed and comprised over 500 million tonnes in total [17]. More recent global estimates are based on acoustic surveys (9 to 15 Gt [18,19]) and modelling studies (0.73 to 16 Gt [20,21,22]). The large variations in estimates between methods are due to a number of factors, including underestimation due to net avoidance, assumptions and parameter choice in models, geographic coverage and extrapolation, and interpretation of acoustics data, for example, whether animal groups, such as siphonophores, are included from the fish fraction [19]. Irrespective of the variation in estimates, the total standing stock of mesopelagic fishes, including myctophids, appears considerably larger than any other commercial fish stock.
Areas of concentration of myctophids in regional estimates from trawl catches, for example, in the western Arabian Sea (100 million tonnes [161]) and the Southern Ocean (containing 70–200 million tonnes [151]), indicate there may be prospective locations in which commercial fisheries for myctophids could be developed. There has been interest in and development of commercial mesopelagic fisheries over the last few decades, but these attempts have, however, not yielded economically viable outcomes in the long-term [162,163]. The former Soviet Union started commercially fishing myctophid fishes (predominantly Diaphus coeruleus and Gymnoscopelus nicholsi) in the Southwest Indian Ocean and Southern Atlantic in 1977, and catches peaked in 1992 with a total harvest of 51,680 tonnes [164]. However, the fishery was eventually abandoned due to low profits [165]. The shelf-edge myctophid Lampanyctodes hectoris was targeted by a purse seine fishery in South African waters with a peak annual catch of 42,560 tonnes in 1973, but the fishery discontinued in the mid-1980s due to the low efficiency of catch [166]. During the 1980s, a rigorous investigation was carried out to assess the possibility of developing a sustainable commercial mesopelagic fishing sector in the northern Arabian Sea [131,167]. Similar trends have been reported for Icelandic fisheries [168] targeting the small hatchetfish, Maurolicus muelleri. Substantial quantities (46,000 and 18,000 tonnes) were caught by pelagic trawling operations in 2009 and 2010, respectively, but the fishery was discontinued because alternative target species were preferred for economic reasons [164].
In more recent years, as the demand for and price of LC omega-3 oils has grown, there has been renewed interest in commercially fishing mesopelagic resources, particularly in nations with strong aquaculture industries, such as Norway [164,169] and Peru [163]. Despite this, the commercial exploitation of mesopelagic fish species continues to face challenges in achieving long-term economic viability [168,170].

4. Future Prospects and Conclusions

Future food requirements for the increasing human population will include additional and alternative fish resources to meet nutritional demands via direct human consumption of fish and indirectly through fish products fed to other domesticated animals. The estimated biomass of myctophids represents a very large resource relative to other fishery stocks and therefore has the potential to contribute to this need. Our review found that most myctophid species have abundant protein and LC omega-3 oil content, along with low levels of contaminants. Several species were found to have very desirable lipid profiles, with many high in DHA, though some species contain large amounts of less desirable wax esters that may impact their practical utilisation. Myctophids also seem to contain good levels of total protein high in arginine and leucine amino acids, in addition to essential trace elements and macro minerals. These findings emphasise the potential of myctophids to provide a valuable source of nutrition for human consumption and to assist agriculture (including aquaculture) production. Our review of the main nutritional components of myctophids, matched with other data, including the price of fish oils and fishmeal, should assist any future assessments of the economic feasibility of harvesting these fishes as an alternative resource. We also demonstrated some major gaps in the knowledge of the nutritional composition of myctophids: a general lack of measurements on the composition of minerals and vitamins in most species, and geographical gaps, including a lack of any protein or contaminant data for myctophids in the Indian Ocean.
Despite the nutritional suitability of myctophids for human consumption, there are barriers to developing sustainable fisheries for them due to major gaps in knowledge regarding their stock assessment and ecology. Estimates of myctophid stock size have an order-of-magnitude variation [164]. Their trophic pathways, links to primary production and roles in biogeochemical cycling are largely unknown [162] despite their being forage species to many high-order fishes, birds and marine mammal predators. Although there is increased demand for LC omega-3 oils, efficient and cost-effective commercial harvesting and processing methods are required for potential myctophid fisheries to attain commercial viability.

Author Contributions

Conceptualization, B.Z. and H.P.; data acquisition: B.Z.; writing—original draft preparation, B.Z., P.D.N., P.V., K.L.-C., H.P. and K.S.; writing—review and editing, B.Z., H.P., P.D.N., P.V., A.W., K.L.-C. and K.S. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Institutional Review Board Statement

Not applicable.

Informed Consent Statement

Not applicable.

Data Availability Statement

Not applicable.

Acknowledgments

B. Zhang acknowledges a PhD scholarship provided by a China Scholarship Council (CSC) Grant from the Ministry of Education of the People’s Republic of China and a University of Tasmania tuition fee scholarship. We are thankful for contributions from the Commonwealth Scientific and Industrial Research Organisation (CSIRO) ResearchPlus Postgraduate Top-Up Scholarship Grants, CSIRO Future Protein Mission and the Holsworth Wildlife Research Endowment. We also acknowledge the late Nansy Phleger for her original watercolour myctophid painting gifted to P.D.N. Four anonymous reviewers and the handling editor Theodora Felegean are thanked for their comments which assisted in improving the manuscript.

Conflicts of Interest

The authors declare no conflict of interest.

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Figure 1. A common myctophid species found in the Southern Ocean is Electrona antarctica (from Nansy Phleger).
Figure 1. A common myctophid species found in the Southern Ocean is Electrona antarctica (from Nansy Phleger).
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Figure 2. Amino acid composition (%, g/100 g of total amino acid, wet weight) of 13 myctophid species caught from the Indian Ocean [25,62,64,65] and trophic Pacific Ocean [63] and Northern Atlantic Ocean [50].
Figure 2. Amino acid composition (%, g/100 g of total amino acid, wet weight) of 13 myctophid species caught from the Indian Ocean [25,62,64,65] and trophic Pacific Ocean [63] and Northern Atlantic Ocean [50].
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Zhang, B.; Pethybridge, H.; Virtue, P.; Nichols, P.D.; Swadling, K.; Williams, A.; Lee-Chang, K. Evaluating Alternative and Sustainable Food Resources: A Review of the Nutritional Composition of Myctophid Fishes. Sustainability 2023, 15, 12039. https://doi.org/10.3390/su151512039

AMA Style

Zhang B, Pethybridge H, Virtue P, Nichols PD, Swadling K, Williams A, Lee-Chang K. Evaluating Alternative and Sustainable Food Resources: A Review of the Nutritional Composition of Myctophid Fishes. Sustainability. 2023; 15(15):12039. https://doi.org/10.3390/su151512039

Chicago/Turabian Style

Zhang, Bowen, Heidi Pethybridge, Patti Virtue, Peter D. Nichols, Kerrie Swadling, Alan Williams, and Kim Lee-Chang. 2023. "Evaluating Alternative and Sustainable Food Resources: A Review of the Nutritional Composition of Myctophid Fishes" Sustainability 15, no. 15: 12039. https://doi.org/10.3390/su151512039

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