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Effects of various organic substrates on growth performance and

nutrient composition of black soldier fly larvae: A meta-analysis

Eko Lela Fitriana, Erika Budiarti Laconi, Dewi Apri Astuti,

Anuraga Jayanegara

PII: S2589-014X(22)00118-9
DOI: https://doi.org/10.1016/j.biteb.2022.101061
Reference: BITEB 101061

To appear in: Bioresource Technology Reports

Received date: 27 January 2022

Revised date: 4 April 2022
Accepted date: 17 April 2022

Please cite this article as: E.L. Fitriana, E.B. Laconi, D.A. Astuti, et al., Effects of various
organic substrates on growth performance and nutrient composition of black soldier fly
larvae: A meta-analysis, Bioresource Technology Reports (2021), https://doi.org/10.1016/
j.biteb.2022.101061

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© 2022 Published by Elsevier Ltd.

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Effects of Various Organic Substrates on Growth Performance and Nutrient

Composition of Black Soldier Fly Larvae: A Meta-analysis

Eko Lela Fitrianaa,c, Erika Budiarti Laconib, Dewi Apri Astutib, Anuraga Jayanegarab,c*

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Graduate School of Nutrition and Feed Science, IPB University, Bogor 16680, Indonesia

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Department of Nutrition and Feed Technology, Faculty of Animal Science, IPB

University, Bogor 16680, Indonesia

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c
Animal Feed and Nutrition Modelling (AFENUE) Research Group, Department of
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Nutrition and Feed Technology, Faculty of Animal Science, IPB University, Bogor 16680,
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Indonesia
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*Corresponding author: Anuraga Jayanegara; Department of Nutrition and Feed

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Technology, Faculty of Animal Science, IPB University, Jl. Agatis Kampus IPB Dramaga,

Bogor 16680, Indonesia. E-mail address: anuraga.jayanegara@gmail.com

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Abstract

This study aimed to evaluate the effects of various organic substrates on the performance

and nutrients of black soldier fly larvae through a meta-analysis. A database was developed

from published articles regarding black soldier fly larvae substrates. The substrates were

categorized into four groups, i.e., animal feed, food waste, manure, and other substrates.

The results showed that larvae reared on animal feed developed at the shortest time, 18

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days, among the substrates (p<0.05). The animal feed substrate also resulted in the highest

poly-unsaturated fatty acid content in larvae 18.81% DM, particularly the C18:2n6 19.93%

DM and C18:3n3 1.82% DM. The larvae’ amino acid and mineral profiles were similar

among the substrates. In conclusion, the response of various substrates to the performance

and nutrient of larvae varied, animal feed substrate had the best performance and nutrient

profiles among the substrates.

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Keywords: bioconversion, energy source, feed, insect, maggot, protein source

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List of abbreviations: -p
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BSFL : black soldier fly larvae
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AF : animal feed

FW : food waste
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MN : manure
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OS : other substrate
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WRR : waste reduction rate

WRI : waste reduction index

CR : conversion rate

FCR : feed conversion rate

GR : growth rate

FLW : fresh larvae weight

DLW : dry larvae weight

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PY : prepupal yield

SR : survival rate

DM : dry matter

CP : crude protein

CF : crude fiber

EPA : eicosapentaenoic acid

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ƩSFA : saturated fatty acid

ƩUFA : unsaturated fatty acid

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ƩMUFA : mono-unsaturated fatty acid

ƩPUFA : poly-unsaturated fatty acid -p

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ƩMCFA : medium-chain fatty acid
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1. Introduction

By 2050, The demand for animal protein is estimated to increase to 25-70% above due to the

rapid growth of the human population, influencing the rising cost of feed (El‐Hack et al., 2020;

Liland 2021). Many of the commonly used feedstuffs, such as maize and soybean meal, may be

affected by this demand. Consequently, finding alternative energy and protein feed sources for

animals is essential. Insects have been considered as potential feed resources, especially as

protein sources. A number of global and local companies have produced insects as feed

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resources (Kawasaki et al., 2019). The utilization of insects as feed has been approved in several

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animals, such as fish and poultry, but their use for ruminants is still debatable (Attivi et al., 2020;

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Bejaei and Cheng, 2020; Bruni et al., 2020; Jayanegara et al., 2017). One of the most exciting
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insects is black soldier fly larvae (BSFL; Hermetia illucens, Diptera: Stratiomydae). BSFL has

been advocated as a protein source rich in essential amino acids, energy (fats), vitamins, and
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minerals (Nekrasov et al., 2019; Shumo et al., 2019; Spranghers et al., 2017). Also, BSFL did not
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contain pathogen factors (Popa and Green, 2012) and produced antimicrobial peptides

(Harlystiarini et al., 2019). From previous studies, the inclusion of BSFL into aquaculture feed
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contributes a particular benefit to the growth performance (Kroeckel et al., 2012). Moreover,
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defatted BSFL biomass was incorporated into broiler feed showed higher digestibility by the

chicken (Schiavone et al., 2017).

More industrialized farming of insects for feed purposes started in Western countries and

started developing in Asian countries. Since then, the interest in insect cultivation as feed has

increased exponentially. In addition, the availability of adequate growing substrate is essential

for sustainable BSFL production. Many studies have evaluated several organic wastes and

formulated substrates used as growing substrates for BSFL (Danieli et al., 2019; El‐Hack et al.,

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2020; Surendra et al., 2020). Organic wastes are superior to other growth substrates because their

reliability is vast and affordable. Using organic waste as a substrate of BSFL contributes to

managing agricultural by-products or waste pollution. Food wastes quantity is reported as more

than half of the municipal solid waste in developing countries (Chhandama et al., 2022).

Further, high quantities of organic waste are a problem and issue related to the environmental

(Silva et al., 2021). On the other hand, organic biomass waste represents the promising eco-

friendly source of sericulture of silkworms (Jaiswal et al., 2021). It means that organic waste is

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the potential to substrate BSFL to produce protein and energy sources and the leftover organic

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waste could be fertilizer (zero-waste management). Various organic wastes that had been

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evaluated as rearing substrates for BSFL were, for instance, almond by-product, beer by-product,
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chicken manure, agri-food leftover, and others (Barbi et al., 2020; Bortolini et al., 2020; Chia et

al., 2020; Palma et al., 2020). Interestingly, BSFL has superior adaptability on poor or
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unbalanced nutrient substrates (Bonelli et al., 2020). Despite its adaptive capability, the
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production and nutrient composition of BSFL are heavily affected by the quality of the substrates

(Ewald et al., 2020; Galassi et al., 2021; Meneguz et al., 2018b).

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Although many studies have reported the effects of various organic substrates and BSFL,
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there is no study to date attempting to summarize those research findings quantitatively.

Therefore, there is a need for an evaluation concerning the effect of various substrates on growth

performance and nutrient composition of BSFL by employing a meta-analysis approach. Meta-

analysis integrates data and statistically analyzes results from articles addressing similar research

and producing general interpretations (Liland et al., 2021). The quantitative evaluation of input

(kind of organic substrate) and output (growth performance, nutrient composition of BSLF) by

the mixed model statistical approach might allow for assessing their relationship. This study,

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therefore, aimed to evaluate the effects of various organic substrates on the growth performance

and the nutrient composition of BSFL through a meta-analysis.

2. Materials and Methods

2.1. Literature search and selection criteria

A database was developed from published articles about various substrates for rearing

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BSFL. The articles were obtained from several electronic databases such as Scopus, Science

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Direct, and Google Scholar, and the keywords used were 'substrate' and 'black soldier fly'. The

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articles were strictly selected by following the Preferred Reporting Items for Systematic Reviews

and Meta-Analyses (PRISMA) protocol. Inclusion criteria of the articles were: (a) published in
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English; (b) various kinds of substrates were reported; (c) reported the performance and/or
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nutrient composition (macronutrient, amino acid profiles, fatty acid profiles, and mineral) of
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BSFL; (d) reported feeding rate of rearing BSFL; and (e) reported replication and variance

(standard deviation, SD or standard error or means, SEM).

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The process of selection is provided in Fig. 1. Briefly, 490 articles were initially identified
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based on the title of articles. In the subsequent step, 108 articles were selected, while 382 were

excluded for several reasons (not published in English, non-appropriate research title, conference

proceedings or review articles). After screening the abstract, we further excluded 68 articles

because of: (i) not relevant contents or variables; (ii) another inclusion such as oil, algae,

microbes, biochar, gypsum, polyethylene, polystyrene, NaCl; (iii) formulated substrate; (iv) pre-

treatment of the substrate (fermentation); (v) without replication of analysis. After evaluating the

full text, we also excluded 9 articles for the following reasons: (i) data in as fed; (ii) incomplete

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information in the method. Finally, 31 articles (122 experiments) were integrated into the

database (Table 1).

2.2. Development of the database

The bibliography, substrate, time of rearing, number of larvae, and source of response

variables were entered in the Microsoft Excel spreadsheet. Sources of heterogeneity included in

the dataset were the feeding system and feeding frequency. Response variables included in the

database consisted of two groups, i.e., growth performance-related parameters and nutrient

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composition-related parameters. The former parameters consisted of waste reduction rate

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(WRR), waste reduction index (WRI), conversion rate (CR), feed conversion rate (FCR), growth

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rate (GR), fresh larva weight (FLW), dry larvae weight (DLW), prepupal yield (PY), and
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survival rate (SR), whereas the latter parameters consisted of chemical macro-nutrient
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composition (dry matter, ash, crude protein, crude fat contents), amino acid profiles, fatty acid

profiles, mineral composition of the BSF. All variables were converted into the same units of
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measurements. The next stage was substrate categorization, in which the substrates were
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categorized into four groups, i.e., food waste (FW), animal feed (AF), manure (MN), and other

substrates (OS; those which could not be categorized into the previous three groups). Descriptive
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statistics of the database are presented in Table 2.

2.3. Statistical analysis

All parameters were checked for the outlier values by employing the boxplot method. After

excluding the outlier values, the data were statistically analyzed using the mixed model meta-

analysis, considering the groups of substrates as fixed effects and the different studies as random

effects. The following mathematical model was used:

Yij = μ + Si + τj + Sτij + eij

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Where Yij = the predicted output for predictor variable Y; μ = mean of the treatment; Si = value of

random effect of study i; τj = fixed effect of the j level; Sτij= random effect between i study and j

level; and eij= residual error. A significant effect was declared at p ≤ 0.05. Tukey’s HSD test

was performed to compare least-square means among the substrate groups. The statistical

analysis employed the PROC MIXED procedure of SAS software (SAS OnDemand for

Academics, online).

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3. Results

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3.1.Various substrates affect BSFL growth performance

The different substrates have different rearing times (Table 3). Larvae reared on AF
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developed at the shortest time among the substrates (p<0.05). The FW substrate had a higher
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WRR value than AF, MN, and OS (p<0.05). The AF and FW substrates had a WRI value more
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elevated than OS. The various substrates significantly impacted different DLW of larvae, where

the AF substrate had the DLW of larvae higher than FW and MN. The various substrate affected
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the SR of larvae; the OS had the lowest value of SR. The CR, FCR, GR, and FLW parameters
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were not affected by various substrates.

3.2.Various substrate affects the nutrient composition of BSFL

The various substrate was associated with ash, crude protein (CP) and crude fat (CF) content

of larvae (Table 4). Ash content in OS was higher than FW and AF substrates (p<0.05). The

larvae reared on the OS had a CP similar to AF but higher than FW substrate (p<0.05). The CF

content was higher for larvae grown on FW than OS but still similar with AF substrate (p<0.05).

The various substrates were not associated with a dry matter (p>0.05). In the underlying order,

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there was a trend for the non-essential amino acids to have higher values in apparently random

data than the essential amino acids (Table 5). The various substrates were not associated with

essential and non-essential amino acid profiles (p>0.05). The lauric acid (C12:00) was the most

abundant in BSFL in all kinds of substrate, followed by palmitic acid (C16:00), linolenic acid

(C18:2n6), and oleic acid (C18:1cis9) (Table 5). The various substrate was associated with a

level of palmitoleic acid (C16:1), trans vaccenic acid (C18:1n7), C18:2n6, α-linolenic acid

(C18:3n3), eicosapentaenoic acid (EPA), total saturated fatty acid (ƩSFA), and total poly-

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unsaturated fatty acid (ƩPUFA) of larvae (p<0.05). The MN has a higher level of C16:1 than AF

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and FW substrates but is still similar to OS (p<0.05). Larvae reared on OS had the highest

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C18:1n7 and EPA value (p<0.05). Larvae on FW substrates had the highest ƩSFA (p<0.05).

Meanwhile, the larvae on AF substrate had the highest ƩPUFA that dominantly with C18:2n6
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and C18:3n3. The larvae reared in all types of substrates contain calcium (Ca), phosphor (P),
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magnesium (Mg), potassium (K), sodium (Na), copper (Cu), iron (Fe), manganese (Mn), and
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zinc (Zn) (Table 6). Potassium was the most dominant mineral in BSFL on all substrate groups,

followed by Ca, Mg, and Fe, while the level of Cu was lower than that of the other minerals. The
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effect of various substrates was not influencing all mineral content of larvae (p>0.05).
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4. Discussion

Utilizing BSFL to transform organic waste can achieve a double purpose in the environment

to resolve the problem of organic waste accumulation and economical feed as a protein source.

Numerous studies have demonstrated that BSFL can grow on various organic substrate sources

such as food waste, agroindustry by-products, animal waste, meat-based products (Surendra et

al., 2020). The physiological and morphological of the midgut BSFL enables BSFL to digest

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these various substrates. BSFL has three parts with different luminal pH in the midgut that plays

a role in enzyme activity to digest substrate and influence gut microbiota. There are; anterior

midgut (pH=6), middle midgut (pH=2), and posterior midgut (pH=8.5) (Bonelli et al., 2019). The

BSFL can modify and digest the substrates properly within four days (Meneguz et al., 2018a).

This study investigated how various substrates would be affecting BSFL performance

parameters. The substrate composition may significantly affect BSFL growth, survival, and

substrate conversion rate. The OS in the present study consists of wastes containing high fiber

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content, dominantly; fresh mussels, rotten mussels, palm decanter, rice straw, fresh mushroom

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root, fermented maize straw, spent coffee, cassava peels, semi-digested grass, and sewage

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sludge. Two (semi-digested grass and sewage sludge) have an extracellular polymeric substance
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presented by bacteria (Leong et al., 2016). Meanwhile, the MN consists of many manures,
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including cattle manure containing material reduction and a long-chain fiber component. The

cattle manure contains high parts of cellulose and lignin contents and commonly lower parts of
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readily available carbon (Lalander et al., 2019). Some of the chemical content of substrates that
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have been mentioned above would be made BSFL digest substrate difficultly and implicate with

long rearing times to result in larvae biomass (Rehman et al., 2019; Oonincx et al., 2015).
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Moreover, the whole organic substrate is more difficult for BSFL to digest than mass or liquid

organic substrate (Surendra et al., 2020). The OS has various particle sizes and complex

compositions in the current study. The OS has the lowest WRR value and is followed by the

lowest WRI value.

The AF in the present study has a good and/or balance in the nutrient composition among

the substrate. It is the reason why the AF has the highest DLW value. Interestingly, the high

value of DLW was implicated with the high value of WRI. It was implied that the substrate is

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being degraded effectively and highly consumed, and absorbed into BSFL biomass. The nutrient

content ratio affected the growth performance and maturity of BSFL. The best ratio for the

substrate nutrient is 1:1 for protein and non-fiber carbohydrate (NFC) (Cammack and Tomberlin,

2017). Furthermore, the variation in substrate degradation rate is not only because of substrate

type but also BSFL strain, feeding rate, and larvae density (Surendra et al., 2020).

On the other hand, the use of AF for BSFL is not economic among FW, MN, and OS. The

AF substrate would be more economical if it is a rejected feed because of the low quality of the

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standard or leftover feed. Another reason is the lack of feed protein sources than an energy

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source. Concerning the economic factor, many researchers were interested in fermentation or

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pretreatment of the substrate to optimize growth performance and nutrient (protein or lipid) of
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BSFL. The addition of 1 wt % of yeast powder into substrate increased the lipid content of BSFL
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and 0.02 wt % yeast would be increased the protein content of BSFL (Wong et al., 2020a; Wong

et al., 2020b). It implied that fermentation might result in an improved outcome. Even though
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fermentation has a positive effect, the result depends on the method of fermentation. The ex-situ
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fermentation method significantly promoted the growth of BSFL more than the in-situ

fermentation method (Wong et al., 2021).

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The OS consists of matter with high fiber content and undigested nutrient composition. The

complex component of OS would be implicated in a low SR value. The BSFL can partially

digest non-detergent fiber (NDF) and an acid detergent fiber (ADF) (Galassi et al., 2021) since

they have a microbe that produces lignocellulose enzymes (Bonelli et al., 2020). However, their

utilization of the NDF and ADF was limited (Galassi et al., 2021).

The study confirmed that the various substrates affected the ash, CP, and CF. Since larvae

degraded ash, it does not build a larvae biomass (Kuttiyatveetil et al., 2019). The high ash level

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in the larvae was due to the time of rearing. A more extended time of rearing would be more ash

content because of the development of their exoskeleton. The exoskeleton of BSFL consists of a

calcium carbonate (CaCO3) which may lead to high calcium and ash content (Baragan-Fonseca

et al., 2017; Shumo et al., 2019). Ash content of larvae was increased by 0-29 days due to them

developing exoskeleton (Do et al., 2020; Hoc et al., 2020).

The CP content of larvae in our study was around 38%-46%. These results are within the crude

protein range for BSFL literature (Chia et al., 2020). The larvae reared on FW had the lowest CP

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would be indicated a CP content of FW substrate. Further research on CP was reported that if

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substrates are unbalanced, the incorporation of crude protein from substrate did not optimum

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(Galassi et al., 2021). In this study, the crude protein content of BSFL was affected by the
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various substrates because several studies in the database were computed from N using a
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conversion factor equal to 6.25, and the other studies were calculated from N equivalent to 4.76.

The differences in conversion factors because of non-protein nitrogen in insects such as nucleic
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acid, phospholipid, chitin and excretion products in the intestinal tract affect the actual protein
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content (Galassi et al., 2021; Janssen et al., 2017; Shumo et al., 2019). Hence, characterizing

both protein amino acid and non-protein nitrogen was necessary.

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The CF of larvae was affected by the CF of the substrate (Palma et al., 2019). Increasing CF

was due to excess protein and non-fiber carbohydrate (NFC) in a substrate (Barragan-Fonseca et

al., 2019; Taufek et al., 2021). There is a high correlation between the level of CF in the larvae

and the NFC of a substrate (Spranghers et al., 2017). Non-fiber carbohydrates from substrate

form several fatty acids that synthesize fat through glycolysis, oxidative decarboxylation, and

then fatty acid synthesis cycles (Ewald et al., 2020; Hoc et al., 2020).

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The amino acid compositions show the quality of protein content. The essential amino acid

of BSFL is comparable with soybean meal and fishmeal (Heuel et al., 2021). In this study,

amino acid compositions did not affect by the various substrate. It supported previous studies

that BSFL had excellent potential as a high-quality feed because it can upgrade waste material

but does not seem to influence the protein content from amino acids (Pinotti et al., 2019; Van

Huis, 2020).

The level of fatty acid of BSFL was influenced by the partially fatty acid of the rearing

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substrate (Ewald et al., 2020). In all substrates in our study, SFA was the highest kind of fatty

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acid in all BSFL. Several SFA as C12:00, myristic acid (C14:00), and 16:00 could be

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synthesized by the other component of the substrate (Ewald et al., 2020; Galassi et al., 2021;
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Spranghers et al., 2017). PUFA would synthesize the SFA such as C10:00, C12:00, C14:00, and
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C16:00 as C18:1cis9 and C18:1n3 via β-oxidation reaction acetyl Co-A (Hoc et al., 2020). The

other explanation of the mechanism is that the NFC, such as glucose and starch content in the
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substrate, could be transformed to fatty acid content in BSFL biomass by glycolysis, acetyl-CoA
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carboxylase, and then fatty acid synthesis pathway (Ewald et al., 2020). It was implied that

larvae properly accumulate PUFA and NFC content of the substrate, but they are finally
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metabolized to synthesize SFA of their body content. Furthermore, C16:00 could be transformed

into C18:00 by fatty acid elongase reaction, either the accumulation from substrates or

biosynthesis in the larvae body.

The present enzymes such as Δ9 desaturase and Δ9 and Δ12fat2 desaturase or Δ6 fat 3

desaturase (Δ12 Δ9 and Δ12 isomerase) in BSFL allows the isomeric change from C18:00 to be

C18:1cis9 and then C18:2n6 and C18:3n3 (Hoc et al., 2020). It could be assumed that the low

level of SFA could be happening because most of them were transformed to be C18:2n6. The

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total of C18:1n7 was accumulated from C18:1n7 in a substrate. In our study, the substrate

influenced the palmitoleic acid (C16:1). The palmitic acid could synthesize C16:1 via isomeric

change with the presence of Δ9 isomerase. The differences in level C16:1 of the BSFL are

probably due to complex factors; level C16:1 on the substrates, the level of NFC on the

substrates, and the presence of various key enzymes that influenced fatty acid synthase in BSFL

(thioesterase II, Δ-9-desaturase, Δ12 fat2 desaturase, or Δ6 fat3 desaturase) (Hoc et al., 2020).

Based on this explanation, the fatty acid profiles of BSFL were influenced by a level of fatty

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acid, NFC, the presence of enzymes that support the mechanism, and the combination of each

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factor.

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BSFL rearing on OS has the highest EPA value due to the accumulation of EPA originating
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from the substrate (Ewald et al., 2020; Liland et al., 2017). The negative trend has been found
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that the decrease EPA is most because lauric acid and SFA increase. (Ewald et al., 2020; Liland

et al., 2017). It was explaining the FW substrate had the highest value of lauric acid (C12:00) or
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SFA and the lowest value of EPA.

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The PUFA is significantly affected by the rearing substrate. It was probably due to higher
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PUFA content in BSFL. Nonetheless, BSFL cannot synthesize PUFA (Liland et al., 2017). The

AF and FW substrate have a high concentration of Unsaturated Fatty Acid (UFA) (Saadoun et

al., 2020). Moreover, the AF substrate consisted of maize distiller, which had high

concentrations of PUFA (Świa̧tkiewicz and Koreleski, 2008).

Mineral has some functions supporting the body to optimize metabolism, such as structural,

physiological, catalytical, and regulatory (Chia et al., 2020). Overall, the mineral levels in BSFL

comply with poultry, pig, and fish requirement (Van Huis, 2020). Mineral K, Ca, Mg, and Fe

were the predominant macro minerals in BSFL biomass in this study. Functionally, Ca is a

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crucial mineral to bone formation and eggshell formation, so deficient or excess of Ca minerals

in feed affect the utilization of the other Ca source. Larvae reared on AF are often inadequate for

this mineral. BSFL could secrete a calcium carbonate deposit (CaCO3) which may account for

the high Ca and ash content (Barragan-Fonseca et al., 2017; Johannsen, 1992; Newton et al.,

1977). The previous study reported a high Ca in BSFL, which the composition exoskeleton of

BSFL might partly explain. Another mineral probably needed in the feed is phosphor (P). based

on the results, the substrate did not influence the P content of larvae.

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5. Conclusion
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Meta-analysis is required for statistical evaluation because the various data about growth
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performance and nutrients of BSFL had been published. The study summarizes 122 research
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findings of the effect of various substrates on the performance and nutrients of BSFL. The results

suggest that all types of substrates influence the performance and nutrients of BSFL. Animal
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feed substrate showed a more effective substrate among the substrates. The presence of poly-
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unsaturated fatty acid, particularly the C18:2n6 and C18:3n3 in high concentration, are
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recommended as substrate considering their positive effect on high poly-unsaturated fatty acid in

BSFL.

Credit authorship contribution statement

Eko Lela Fitriana: Data collection, investigation, data input, writing the original draft, formal

analysis, figure visualization

Erika Budiarti Laconi: Supervision, manuscript correction

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Dewi Apri Astuti: Supervision, manuscript correction

Anuraga Jayanegara: Supervision, conceptualization, reference collection, data curation,

writing/reviewing the manuscript

Declaration of competing interest

The authors declare that they have no competing interests.

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Funding: This work was supported by Indonesian Ministry of Education within the Scheme of

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Penelitian Pendidikan Magister menuju Doktor untuk Sarjana Unggul (PMDSU) with contract

200/SP2H/PMDSU/DRPM/2020.
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Table 1. Studies included in the meta-analysis

Feeding Rate
Reference Study Exp. Substrate Feeding System Rearing time (d)
(g/larvae/day)

(Myers et al., 2008) 1 1-4 Dairy manure Daily 30 0.003

Dairy manure Daily 28 0.005
Dairy manure Daily 26 0.007

(Diener et al., 2009) 2 5-9

Dairy manure
Chicken feed
Chicken feed
Daily

o
Interval (2-3 days)
Interval (2-3 days)
f 26
42
33
0.009
0.013
0.025
Chicken feed
r o
Interval (2-3 days) 20 0.050
Chicken feed
Chicken feed
- p
Interval (2-3 days)
Interval (2-3 days)
17
16
0.100
0.200
(Tschirner and Simon, 2015) 3 10-12 Animal feed
DDGS
Dried sugar beet
re Batch
Batch
Batch
15
15
15
0.025
0.022
0.022
(Leong et al., 2016) 4 13-21 Sewage sludge

l
Sewage sludge P Daily
Daily
8
8
0.005
0.025

n a
Sewage sludge
Fruit waste
Daily
Daily
8
10
0.125
0.005

u r Fruit waste
Fruit waste
Daily
Daily
10
10
0.025
0.125

(Manurung et al., 2016) 5 Jo 22-26

Palm Decanter
Palm Decanter
Palm Decanter
Rice straw
Daily
Daily
Daily
Daily
13
13
13
54
0.005
0.025
0.125
0.013
Rice straw Daily 54 0.025
Rice straw Daily 53 0.050
Rice straw Daily 41 0.100
Rice straw Daily 38 0.200
(Supriyatna et al., 2016) 6 27-31 Cassava peel Interval (3 days) 54 0.013
Cassava peel Interval (3 days) 42 0.025
Cassava peel Interval (3 days) 29 0.050
Cassava peel Interval (3 days) 20 0.100
Cassava peel Interval (3 days) 20 0.200

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(Spranghers et al., 2017) 7 32-35 Chicken feed Interval (3 days) 12 0.050
Digested Interval (3 days) 15 0.040
Vegetable waste Interval (3 days) 15 0.040
Restaurant food waste Interval (3 days) 18 0.033
(Jucker et al., 2017) 8 36-38 Fruits’ waste Batch 52 Ad libitum
Vegetable waste Batch 48 Ad libitum
Fruit and vegetable waste Batch 36 Ad libitum
(Nyakeri et al., 2017) 9 39-42 Banana peelings Interval (4 days) 16 0.100
Brewer’s waste Interval (4 days) 16 0.100
Human fecal sludge
Restaurant food waste
Interval (4 days)
Interval (4 days)
o f 16
16
0.100
0.100
(Rehman et al., 2017) 10 43-44 Dairy manure
Soybean curd
Batch

r
Batch
o 24
19
0.042
0.053
(Liu et al., 2018) 11 45-47 Brewer’s waste
Pig manure
- p
Interval (5 days)
Interval (5 days)
15
17
0.200
0.200

(Meneguz et al., 2018b) 12 48-51

Semi-digested grass
Fruits & veg waste
r e Interval (5 days)
Interval
70
20
0.200
Ad libitum
Fruit’s waste

l P
Winery by-product
Brewery by-product
Interval
Interval
Interval
22
22
8
Ad libitum
Ad libitum
Ad libitum
(Kinasih et al., 2018) 13 52-66

n a
Vegetable waste
Vegetable waste
Interval (3 days)
Interval (3 days)
51
48
0.013
0.025

u r Vegetable waste
Vegetable waste
Interval (3 days)
Interval (3 days)
38
28
0.050
0.100

Jo Vegetable waste
House manure
House manure
House manure
Interval (3 days)
Interval (3 days)
Interval (3 days)
Interval (3 days)
23
49
39
27
0.200
0.013
0.025
0.050
House manure Interval (3 days) 22 0.100
House manure Interval (3 days) 22 0.200
Tofu dreg Interval (3 days) 35 0.013
Tofu dreg Interval (3 days) 33 0.025
Tofu dreg Interval (3 days) 29 0.050
Tofu dreg Interval (3 days) 29 0.100
Tofu dreg Interval (3 days) 27 0.200
(Cai et al., 2019) 14 67 Fresh mushroom roots Batch 20 0.050
(Bava et al., 2019) 15 68-71 Hen Diet Batch 15 Ad libitum
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Maize Distiller Batch 16 Ad libitum
Okara Batch NA Ad libitum
Brewer’s Grains Batch 22 Ad libitum
(Lalander et al., 2019) 16 72-82 Abattoir waste Interval (3 days) 17 0.040
Abattoir-fruits veg Interval (3 days) 17 0.040
Dog food Interval (3 days) 18 0.040
Digested sludge Interval (3 days) 42 0.040
Food waste Interval (3 days) 19 0.040
Fruits & veg Interval (3 days) 47 0.040
Human feces
Poultry feed
Interval (3 days)
Interval (3 days)
o f 19
16
0.040
0.040
Poultry manure
Primary sludge
r o
Interval (3 days)
Interval (3 days)
19
32
0.040
0.040

(Beniers and Graham, 2019) 17 83

Undigested sludge
Chicken feed
- p
Interval (3 days)
Daily
51
11
0.040
0.015
(Lim et al., 2019) 18 84-85 Coconut endosperm

r
Soybean curd residue
e Batch
Batch
15
20
0.033
0.025
(Giannetto et al., 2020a)
(Ewald et al., 2020)
19
20
86
87-91 Bread
Fish
l P
Fruit and vegetable waste Daily
Batch
Batch
NA
14
14
0.075
0.014
0.014

n a
Food waste
Fresh mussels
Batch
Batch
14
14
0.014
0.143

(Giannetto et al., 2020b) 21

u r
92
Rotten mussels
Fruit and vegetable waste
Batch
Daily
14
NA
0.143
0.075
(Jucker et al., 2020)

(Kierończyk et al., 2020)

22

23
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95-98
Cricket waste
Locust waste
Wheat bran
Carrots
Batch
Batch
Batch
Batch
16
18
13
13
Ad libitum
Ad libitum
0.077
0.077
Cabbages Batch 13 0.077
Potatoes Batch 13 0.077
(Saadoun et al., 2020) 24 99-100 Brewery by-products Batch 33 0.061
Tomato peels and seeds Batch 24 0.083
(Scala et al., 2020) 25 101-104 Apple Batch 16 0.044
Banana Batch 14 0.050
Apple + banana Batch 15 0.047
Brewery spent grain Batch 11 0.064
(Adebayo et al., 2021) 26 105-108 Chicken feed Interval (2 days) 21 0.029
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Brewery waste Interval (2 days) 23 0.026
Food remains Interval (2 days) 28 0.021
Fruits’ waste Interval (2 days) 38 0.016
(El-Dakar et al., 2021) 27 109-112 Fresh pig manure Interval (2 days) 36 Ad libitum
Fresh goat manure Interval (2 days) 26 Ad libitum
Fresh quail manure Interval (2 days) 30 Ad libitum
Fresh poultry manure Interval (2 days) na Ad libitum
(Galassi et al., 2021) 28 113-116 Hen diet Batch 15 0.050
Okara Batch 18 0.050
Maize distillers
Brewer’s grains
Batch
Batch
o f 16
22
0.050
0.050
(Klammsteiner et al., 2021) 29 117-118 Chicken feed
Food waste
r o
Interval (2 days)
Interval (2 days)
21
22
0.100
0.170
(Raksasat et al., 2021) 30 119-120 Sewage sludge
Palm kernel meal
- p
Batch
Batch
28
13
0.018
0.038
(Fischer et al., 2021) 31 121-122 Spent coffee
Donut dough
r e Batch
Batch
35
35
0.022
0.022
*NA= not available;

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Table 2. Descriptive statistics of the database used in the meta-analysis

Variable Unit N Mean SD* Min Max

Growth performance
Time of rearing Day 114 23.78 12.627 8 70
WRR* % 57 51.91 22.078 9.290 96.860
WRI* g/day 35 1.90 1.608 -0.070 5.300
CR* 73 0.21 0.586 0.0003 5.0600
FCR* 12 3.11 2.632 0.870 9.290
GR* g/day 29 0.09 0.191 -0.070 0.650
FLW* mg 48 169.83 68.540 34.80 290.00
DLW* mg 22 54.40 43.169 20 160
SR* % 49 86.16 15.218 39.0 101.5
Macro-nutrient composition
DM* % 39 32.40 11.156 8.3 76.6

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Ash % DM 32 10.36 5.470 2.7 22.9
CP* % DM 45 42.72 7.366 30.750 55.000
CF* % DM 38 31.80 6.769 8.610 46.000
Amino acid profile
Arginine % DM
% DM
14
14
pr4.33
5.74
1.702
3.585
0.390
0.710
6.490
16.490
Alanine
e-
Aspartic Acid % DM 14 7.34 3.043 0.770 9.280
Glutamic Acid % DM 14 9.56 4.797 0.960 21.760
Pr

Glycine % DM 14 4.51 1.876 0.650 5.920

Histidine % DM 14 2.88 1.665 0.30 7.490
Isoleucine % DM 14 3.66 1.534 0.50 4.760
Leucine % DM 14 5.72 2.487 0.710 8.010
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Lysine % DM 14 5.31 2.217 0.620 8.250

Methionine % DM 14 1.45 0.689 0.180 2.050
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Phenylalanine % DM 14 3.23 1.391 0.420 4.420

Proline % DM 13 4.50 1.775 0.600 5.870
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Serine % DM 14 3.40 1.347 0.350 4.390

Threonine % DM 14 3.14 1.197 0.380 3.860
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Tryptophan % DM 13 1.36 0.560 0.120 1.790

Valine % DM 14 5.98 2.022 0.750 6.590
Cystine % DM 13 0.52 0.208 0.070 0.670
Tyrosine % DM 14 5.41 2.937 0.570 10.880
Fatty acid profile
C10:00 % DM 16 1.08 0.337 0.550 1.690
C12:00 % DM 28 36.74 12.065 16.610 68.000
C14:00 % DM 27 6.71 2.000 2.850 10.360
C16:00 % DM 29 14.08 4.050 5.780 20.500
C18:00 % DM 29 2.33 1.104 0.720 4.900
C20:00 % DM 8 0.85 1.071 0.017 2.580
C14:1 % DM 11 0.46 0.368 0.040 1.320
C16:1 % DM 28 4.45 3.076 0.280 11.540
C18:1n7 % DM 9 0.81 0.486 0.100 1.500
C18:1cis9 % DM 26 11.18 3.991 4.270 23.980

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C18:1cis 11 % DM 8 0.38 0.105 0.280 0.580

C18:2n6 % DM 26 11.32 7.792 1.370 28.400
C18:3n3 % DM 27 1.28 0.900 0.00 3.40
EPA* % DM 5 0.89 0.992 0.00 2.00
ƩSFA* % DM 28 64.76 10.062 41.300 86.000
ƩUFA* % DM 6 34.23 12.039 24.200 56.800
ƩMUFA* % DM 25 20.33 8.062 10.100 42.57
ƩPUFA* % DM 20 11.46 7.071 1.370 24.10
ƩMCFA* % DM 8 25.02 13.645 4.770 41.700
Mineral composition
Ca % DM 10 1.45 1.122 0.521 3.935
P % DM 8 0.62 0.389 0.176 1.322
Mg % DM 8 1.18 0.908 0.260 2.647
K % DM 8 1.86 0.742 0.790 2.660

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Cu % DM 10 0.045 0.030 0.011 0.090

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Mn % DM 8 0.33 0.194 0.160 0.730
Na % DM 7 0.80 0.825 0.060 2.500
Fe % DM 11 1.15 1.139 0.084 2.970
Zn % DM 11 pr 0.29 0.392 0.040 1.110
*SD= standard of deviation; WRR= waste reduction rate; WRI= waste reduction index; CR= conversion rate; FCR= feed
e-
conversion rate; GR= growth rate; FLW= fresh larvae weight; DLW= dry larvae weight; SR= survival rate; DM= dry
matter; CP= crude protein; CF= crude fat; EPA= eicosapentaenoic acid; SFA= saturated fatty acid; UFA= unsaturated
fatty acid; MUFA= mono-unsaturated fatty acid; PUFA= poly-unsaturated fatty acid; MCFA=medium-chain fatty acid;
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Table 3. Performance Parameters of BSFL

Response variables Unit Animal Feed Food Waste Manure Other SEM* P-value
Time of rearing Day 18±5.8c 24±13.2bc 25±6.4ab 31±18.1a 0.102 <0.001
WRR* % 54.6±17.27b 69.4±15.05a 44.5±14.69bc 30.8±19.20c 1.899 <0.001
WRI* g/day 2.81±1.46a 2.13±1.57a NA 0.42±0.31b 0.005 <0.001
CR* 0.230±0.1134 0.148±0.0907 0.138±0.0460 0.319±1.1180 0.0002 0.484
FCR* 2.67±2.503 3.17±3.326 3.80±0.566 NA 0.037 0.895
GR*
FLW*
g/day
mg
0.077±0.2148
180.2±72.47
0.189±0.2553
174.8±77.62
NA
169.6±51.90
o f 0.031±0.0831
113.6±29.60
0.0002
11.024
0.095
0.204
DLW* mg 76.0±44.99a 23.5±3.11b
r o
23.0±2.47b NA 0.209 0.010
SR* % 91.12±11.857a 88.62±8.922a

- p
93.59±3.441a 72.57±19.924b 0.004 0.043
*SEM= standard error of mean; WRR= waste reduction rate; WRI= waste reduction index; CR= conversion rate; FCR= feed conversion rate; GR= growth rate; FLW= fresh
larvae weight; DLW= dry larvae weight; SR= survival rate; NA= not available;

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Table 4. Macro-chemical Nutrient of BSFL

Response variables Unit Animal Feed Food Waste Manure Other SEM* P-value
DM* % 36.37±11.087 29.12±10.120 26.55±6.576 27.41±11.284 0.126 0.295
Ash % DM 8.32±3.495b 10.40±6.256b NA 20.33±2.026a 0.032 0.006
CP % DM 44.66±7.524ab 38.43±5.626b NA 46.03±6.085a 0.089 0.044
CF % DM 30.72±4.978ab 35.69±5.274a NA 26.04±11.238b 0.002 0.040
*SEM= standard error of mean; DM= dry matter; NA= not available;

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Table 5. Amino acid and fatty acid profiles of BSFL

Response variables Unit Animal Feed Food Waste Manure Other SEM* P-value
Amino acid
Essential
Histidine % DM 3.04±0.420 3.35±2.988 3.20±0.148 2.10±1.300 0.105 0.608
Isoleucine % DM 4.62±0.113 2.67±1.852 4.39±0.403 3.34±1.915 0.055 0.254
Leucine % DM 7.23±0.590 4.13±3.067 6.81±0.276 5.27±3.090 0.132 0.321
Lysine
Phenylalanine
% DM
% DM
6.07±0.277
3.86±0.225
4.42±3.162
2.31±1.700
6.62±0.636
4.31±0.156
o f 4.79±2.785
2.98±1.707
0.150
0.041
0.684
0.261
Threonine
Valine
% DM
% DM
3.69±0.086
6.23±0.092
2.56±1.532
3.91±2.471
r o
3.80±0.049
6.28±0.438
2.75±1.587
4.57±2.576
0.051
0.112
0.355
0.272
Methionine % DM 1.76±0.044 0.95±0.864
-p 1.95±0.141 1.41±0.818 0.007 0.279
Cysteine
Non-essential
% DM 0.60±0.034 0.37±0.257

r e 0.61±0.085 0.51±0.295 0.002 0.129

Arginine
Alanine
% DM
% DM
4.85±0.144
5.69±0.327
l P
4.12±2.525
7.15±6.658
5.01±0.127
6.09±0.622
3.69±2.210
4.22±2.301
0.156
0.465
0.492
0.440
Aspartic Acid
Glutamic Acid
% DM
% DM
8.89±0.395
9.78±0.540
n a 5.54±3.749
10.53±8.547
9.10±0.184
10.13±0.658
6.70±3.961
7.75±4.516
0.256
1.097
0.316
0.473
Glycine
Proline
% DM
% DM
u r
5.49±0.296
5.07±0.578
3.32±2.305
3.67±2.657
5.67±0.361
5.50±0.417
4.16±2.365
4.05±2.333
0.083
0.177
0.304
0.769
Serine
Tryptophan
Tyrosine
% DM
% DM
% DM J o
4.01±0.113
1.55±0.145
6.57±1.379
2.73±1.691
0.97±0.733
3.23±2.926
4.12±0.389
1.66±0.049
8.55±3.302
3.08±1.837
1.33±0.780
4.85±2.864
0.063
0.017
0.089
0.409
0.102
0.169
Fatty Acid
C10:00 % DM 1.00±0.566 1.10±0.251 1.29±0.378 0.71±0.219 0.0001 0.427
C12:00 % DM 31.44±4.028 41.80±12.533 36.15±12.397 30.14±16.144 0.019 0.101
C14:00 % DM 6.18±0.426 7.23±2.284 5.76±1.034 6.98±3.730 0.015 0.092
C16:00 % DM 16.04±3.518 12.23±3.426 14.38±5.761 15.82±3.950 0.004 0.119
C18:00 % DM 2.64±1.006 2.21±1.266 2.06±1.129 2.27±0.903 0.004 0.564
C20:00 % DM 1.30±1.153 0.10±0.000 NA NA 0.108 0.877
C14:1 % DM NA 0.28±0.268 0.73±0.429 0.34±0.247 0.0003 0.406
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C16:1 % DM 3.27±1.658b 3.01±1.631b 10.02±1.048a 6.32±3.798a 0.015 <0.001
C18:1n7 % DM 0.79±0.355ab 0.25±0.212b NA 1.40±0.141a 0.219 0.030
C18:1cis9 % DM 11.80±2.649 9.66±3.171 15.58±6.146 9.95±3.145 0.029 0.177
C18:1cis 11 % DM 0.44±0.140 0.34±0.068 NA NA 0.0005 0.399
C18:2n6 % DM 19.93±5.086a 7.63±3.106b 3.92±4.196b 7.63±7.376b 0.080 <0.001
C18:3n3 % DM 1.82±0.994a 1.38±0.612a 0.06±0.074b 0.94±0.461ab 0.0009 0.017
EPA % DM NA 0.25±0.354b NA 1.31±1.110a 0.012 0.022
% DM 59.52±9.196b 70.98±8.948a 59.66±7.638ab 69.65±5.728ab 0.938 0.009

f
ƩSFA
% DM 35.10±3.394 33.80±15.394 NA NA 3.927 0.643

o
ƩUFA
% DM 23.05±11.285 16.34±4.777 26.27±6.479 22.85±4.738 0.944 0.155

o
ƩMUFA
ƩPUFA
ƩMCFA
% DM
% DM
18.81±4.493a
23.61±4.857
11.59±6.004ab
25.87±17.654
p r
3.98±4.273b
NA
7.50±0.990ab
NA
0.058
26.629
0.032
0.805
*SEM= standard error of mean; NA= not available;

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Table 6. Mineral composition of BSFL
Response variables
Unit Animal Feed Food Waste SEM* P-value
Ca % DM 1.569±1.6170 1.368±0.8225 0.007 0.838
P % DM 0.609±0.3299 0.623±0.4949 0.002 0.117
Mg % DM 1.064±0.8214 1.301±1.1002 0.007 0.894
K % DM 1.754±0.7320 1.958±0.8484 0.003 0.563
Cu
Mn
% DM
% DM
0.024±0.0105
0.358±0.2558
0.059±0.0310
0.293±0.1414
o f 1.524e-6
0.0002
0.233
0.938
Na
Fe
% DM
% DM
0.618±0.3961
1.003±1.3296
r o
1.050±1.2834
1.239±1.1203
0.005
0.008
0.086
0.916
Zn
*SEM= standard error of mean;
% DM 0.140±0.0976

- p
0.377±0.4773 0.003 0.459

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Research publication:
Identificatio
Articles were collected from Scopus, Google
scholar using the keywords ‘substrate’ and
‘black soldier fly’ (n=490) Exclude based on kind of article (a
n

proceeding or review articles), not

published in English, and non-
appropriate research title (n=382)

Exclude based on:

Screening

Review on abstract:
Number of articles based on a review of (i) Not relevant contents and
abstract (n=108) variables (n=15)
(ii) Another includes oil, algae,
microbes, biochar, gypsum,
NaCl, polyethylene,

f
polystyrene. was used (n=19)

oo
(iii) Formulated substrate (n=27)
(iv) Pre-treatment of the substrate
Full-text evaluation:
Eligibility

such as aerobic or anaerobic

Articles for database development (n=40) fermentation (n=6)
pr (v) Without replication
analysis (n=1)
of
e-
Exclude based on:
Pr

(i) Data in as fed (n=1)

Final database: (ii) Incomplete information in the
Included

Number of articles used for meta-analysis method (n=8)

(n=31)
al
rn

Fig 1. Flow charts of publications utilized for the meta-analysis

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Graphical Abstract

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Highlights

 A meta-analysis was performed to evaluate the effects of various organic substrates on

BSF larvae.

 Animal feed substrate had the optimum rearing time and dry weight of the larvae.

 Polyunsaturated fatty acid contents in larvae were affected by type of substrate.

 Amino acid and mineral contents of larvae were not affected by type of substrate.

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