Performance and body composition of Nile tilapia fed diets containing ora-pro-nóbis leaf meal

Abstract

Recent studies have indicated that ora-pro-nóbis leaf meal (OLM) contains high levels of digestible amino acids compared with conventional ingredients such as corn and wheat bran. This study evaluated effects of OLM on productive performance and body chemical composition of juvenile Nile tilapia. Four hundred fish (14.16 ± 0.43 g) were distributed into 20 tanks (1 m³) in a completely randomized design with five treatments (0, 3.5, 7, 10.5, and 14 % inclusion) and four replicates over 58 days. Inclusion of OLM did not affect fish survival and improved productive performance, with higher final weight (86.18 g), weight gain (72.35 g), daily weight gain (1.24 g), and fillet weight (27.97 g) observed at the 14 % inclusion level, with no statistical differences compared with levels from 7 % onward. Feed intake averaged 1.83 g and did not differ among diets containing OLM, although values were higher than control treatment. In carcass composition, ether extract decreased at inclusion levels of 10.5 % and 14 %, with values of 7.08 % and 7.04 %, respectively, whereas crude protein increased from the 7 % inclusion level onward, reaching values of 16.76 %, 16.91 %, and 17.24 % at inclusion levels of 7 %, 10.5 %, and 14 %, respectively. Therefore, OLM demonstrates potential as an alternative ingredient for tilapia diets, contributing to economic sustainability and competitiveness of aquaculture, especially for small-scale producers, while also supporting regional circular economy systems.

Keywords:
alternative feedstuff; weight gain; tilapia farming; visceral fat index.

Resumo

Estudos recentes indicam que a farinha de folhas de ora-pro-nóbis (FFO) apresenta teores elevados de aminoácidos digestíveis em comparação a ingredientes convencionais, como milho e farelo de trigo. Nesse contexto, este estudo avaliou os efeitos da FFO sobre o desempenho produtivo e a composição química corporal de juvenis de tilápia-do-Nilo. Quatrocentos peixes (14,16 ± 0,43 g) foram distribuídos em 20 tanques (1 m³), em delineamento inteiramente casualizado com cinco tratamentos (0; 3,5; 7; 10,5 e 14 % de inclusão) e quatro repetições, durante 58 dias. A inclusão de FFO não afetou a sobrevivência dos peixes e promoveu melhorias no desempenho produtivo, com maiores valores de peso final (86,18 g), ganho de peso (72,35 g), ganho de peso diário (1,24 g) e peso de filé (27,97 g) no nível de 14 %, sem diferenças estatísticas em relação aos níveis a partir de 7 %. O consumo de ração foi de 1,83 g, não diferindo das dietas que continham FFO, embora os valores tenham sido maiores do que no tratamento controle. Na composição de carcaça, observou-se redução do extrato etéreo (7,08 % e 7,04 %) nos níveis de 10,5 % e 14 %, respectivamente, e aumento da proteína bruta a partir de 7 % de inclusão, com valores de 16,76 %, 16,91 % e 17,24 % para os níveis de 7 %, 10,5 % e 14 %, respectivamente. Dessa forma, a FFO demonstra potencial como ingrediente alternativo na alimentação de tilápias, contribuindo para a sustentabilidade econômica e a competitividade da aquicultura, especialmente para pequenos produtores, além de favorecer a economia circular regional.

Palavras-chave:
alimentação alternativa; ganho de peso; tilapicultura; índice de gordura visceral.

1. Introduction

The Food and Agriculture Organization (FAO) of the United Nations (¹) has predicted a significant increase in global aquatic organism production by 2032. Global production is projected to reach 205 million tons, with 111 million tons derived from aquaculture alone, representing a 17 % increase. This growth is expected to directly affect demand for aquatic foods, which is projected to increase by 12 % to meet an estimated average per capita consumption of 21.3 kg.

Under this scenario, adoption of alternative feed ingredients has become a key strategy to support sustainable aquaculture growth. Brazil stands out because of its biodiversity and availability of agro-industrial products and by-products with potential for inclusion in aquafeeds (²). However, studies evaluating these ingredients remain limited despite their importance for improving production efficiency and economic viability (³).

High feed costs remain one of the main constraints in aquaculture, making development of cost-effective nutritional strategies essential (⁴). Among alternative ingredients, leaf meals represent low-cost plant protein sources and may partially replace conventional feedstuffs when properly incorporated into diets (⁵).

Besides reducing costs, alternative ingredients may contribute to regional circular economies, particularly in small-scale and subsistence aquaculture systems. Leaf meals from easily cultivated plants can be locally produced, enabling artisanal feed formulation, improving access to fish as a high-quality protein source, and supporting vulnerable communities (⁶). Ingredient diversification also increases flexibility in feed formulation by allowing partial replacement of traditional inputs such as fish meal, fish oil, corn, and soybean meal according to price fluctuations and ingredient availability (⁷).

Ora-pro-nóbis (Pereskia aculeata) is a plant species belonging to the Cactaceae family that occurs in tropical regions ranging from southern United States to southern Brazil and is distributed throughout North, Central, and South America (⁸). Its adaptability to low-fertility soils and high biomass productivity makes it a promising candidate for feed applications (⁹). Souza et al. (¹⁰) reported yields of up to 25.6 t ha⁻¹ year⁻¹ of dry biomass, corresponding to 5,759 kg ha⁻¹ year⁻¹ of vegetable protein.

Studies on alternative ingredients are essential to determine optimal inclusion levels and their effects on growth performance, feed efficiency, and carcass composition, all of which are directly associated with production profitability. Recent findings have indicated that ora-pro-nóbis leaf meal (OLM) contains higher levels of digestible amino acids than conventional ingredients such as corn and wheat bran (¹¹). Moreover, relevance of such alternatives is reinforced by rapid expansion of tilapia farming in Brazil, with production increasing by 103 % over the last decade, reaching 579 thousand tons in 2023 and accounting for 65.3 % of national farmed fish production (¹²).

Considering the importance of Nile tilapia for both Brazilian and global aquaculture, as it is the second most cultivated fish species worldwide (¹³), alternative feed ingredients have been continuously sought to support sustainable expansion of this activity. Therefore, studies assessing inclusion of ora-pro-nóbis leaf meal (OLM) in juvenile Nile tilapia diets remain limited, particularly regarding its effects on feed use efficiency and nutrient deposition.

Based on the above, this study aimed to evaluate effects of OLM on growth performance and body chemical composition of juvenile Nile tilapia fed diets containing increasing inclusion levels of this ingredient.

2. Material and methods

2.1 Experimental design

Four hundred juvenile Nile tilapia (14.16 ± 0.43 g) were distributed into 20 tanks (1 m³) in a completely randomized design consisting of five treatments and four replicates, with 20 fish per experimental unit (tank), over a 58-day interval. Experimental units were connected to a geomembrane tank (30,000 L) containing biological media and aquatic plants, which functioned as a biofilter in a recirculating system.

The OLM was obtained from ora-pro-nóbis leaves cultivated locally during summer/fall 2022. Leaves were dried in a forced-air oven at 55 °C for 72 h, subsequently ground through a 0.5-mm sieve, and stored under refrigeration in plastic bags. Based on dry matter composition of the OLM used (Table 1) and digestibility values reported previously (¹¹), four experimental diets were formulated (Table 2).

Table 1
Chemical composition of ora-pro-nóbis leaf meal on a dry matter-basis.

		Bromatological Chemical Composition
	DM%	CP% EE% MM% FB% NDF% ADF%	GE (Mj/kg)
OLM	95.56	24.36 4.54 20.60 18.96 42.28 8.91	15.54

Table 2
Formulation and chemical composition of experimental diets containing increasing levels of orapro-nóbis leaf meal on a natural matter basis.

		Inclusion levels (%)
Ingredients	0	3.5	7	10.5	14
Soybean meal	61.67	61.90	61.90	62.01	62.13
Ora-pro-nóbis leaf meal	0.00	3.50	7.00	10.50	14.00
Corn	32.51	27.38	23.08	18.65	13.4
CaHPO4	2.50	2.90	2.90	2.80	3.00
Soy oil	2.50	3.70	4.50	5.42	6.75
Vitamin-mineral mixture 1	0.50	0.50	0.50	0.50	0.50
Salt	0.10	0.10	0.10	0.10	0.10
BHT2	0.01	0.01	0.01	0.01	0.01
CaCO3	0.20	0.00	0.00	0.00	0.10
Adsorbent	0.01	0.01	0.01	0.01	0.01
Composition analyzed (Nutrient) Moisture	3.11	4.26	4.24	4.98	4.88
Crude protein	31.23	31.62	31.99	32.40	32.73
Digestible protein 3	28.00	28.01	28.00	28.02	27.97
Gross energy (MJ/kg)	4047.83	4100.45	4138.19	4186.73	4242.41
Digestible energy (MJ/kg) 3	3002.00	3009.00	3001.00	3002.00	3014.00
Ether extract	4.07	4.81	5.41	7.06	7.94
Mineral matter	9.09	7.77	8.38	8.66	9.37
Starch 4	28.01	24.81	22.13	19.36	16.09
Crude fiber	5.99	5.98	5.34	5.97	6.63

• ¹
  The guaranteed levels of vitamin and mineral supplements per kilogram of product were as follows: vit. A = 1,200,000 IU; vit. D3 = 200,000 IU; vit. E = 12,000 mg; vit. K3 = 2400 mg; vit. B1 = 4800 mg; vit. B2 = 4800 mg; vit. B6 = 4000 mg; vit. B12 = 4800 mg; folic acid = 1200 mg; calcium pantothenate = 12,000 mg; vit. C = 48,000 mg; biotin = 48 mg; choline = 65,000 mg; niacin = 24,000 mg; Fe = 10,000 mg; Cu = 6000 mg; Mn = 4000 mg; Zn = 6000 mg; I = 20 mg; Co = 2 mg; and Se = 20 mg.
• ²
  BHT- butylated hydroxytoluene, NDF- neutral detergent fiber, ADF - acid detergent fiber.
• ³
  Values were calculated using digestibility coefficients determined by (¹¹,¹⁶);
• ⁴
  Values were c alculated using tabulated data (¹⁷).

Isoproteic and isoenergetic diets for digestible protein and digestible energy, respectively (¹⁴), were formulated with inclusion levels of 3.5 %, 7 %, 10.5 %, and 14 % OLM, whereas control diet contained no OLM (Table 2). Inclusion levels of OLM were defined based on digestibility values reported by Matos et al. (¹¹), and nutritional requirements of Nile tilapia described by Carneiro et al. (¹⁵). Ground ingredients were extruded into 2.0-mm pellets using an extruder (Exteec® Machines, SP, Brazil) and dried in a forced-air circulation oven at 55 °C for 48 h. Produced feeds were stored at 8 °C until use.

Feeding management consisted of three daily feeding events until apparent satiety of Nile tilapia (8:30 a.m., 1:30 p.m., and 5:30 p.m.), with feed allowance limited to 1.5 % of average fish biomass in each tank.

2.2 Performance analysis and body yields

At the end of the experimental period, all fish were anesthetized by immersion in a eugenol anesthetic solution diluted in alcohol (5:95) at a ratio of 1 mL per liter of water (¹⁸). Fish were then counted, measured, and weighed to determine average final weight (AFW) and, subsequently, average weight gain (AWG), daily weight gain (DWG), daily feed intake (DFI), feed conversion ratio (FCR), protein efficiency ratio (PER), feed efficiency (FE), and survival rate (SURV).

All 20 fish from each experimental unit were euthanized in an excess eugenol solution (2 mL per liter of water) to evaluate body yield parameters and chemical composition. Of these, seven fish were randomly selected, frozen whole, and used for body chemical composition analyses, whereas another seven fish were randomly selected for evaluation of body weight with head (BWH) and subsequently frozen for chemical analyses. The remaining six fish were used to determine fillet weight (FW) and following indices: hepatosomatic index (HSI), viscerosomatic index (VSI), and visceral fat index (VFI). Fillets were also frozen for subsequent chemical composition analyses.

2.3 Chemical composition analysis

Fillets, carcasses, and whole fish were identified and stored at -20 °C. One day before processing, samples were transferred to a refrigerator for slow thawing. Samples were processed on following day while still partially frozen and with firm tissues, according to Matos et al. (¹⁹), until desired particle size (powder) was obtained.

Crude protein (CP) was determined using micro-Kjeldahl distillation method, whereas moisture (MOI) and mineral matter (MM) contents were determined according to AOAC guidelines (²⁰). Ether extract (EE) was determined using a Goldfish apparatus (Tecnal Ltda.) by hot solvent extraction with petroleum ether (30-70 °C). Gross energy of experimental diets was determined using an adiabatic bomb calorimeter.

2.4 Water quality

Water temperature, dissolved oxygen (DO) concentration, and pH were determined using a digital multiparameter meter (Model AK-88, Akso Ltda.). Water temperature and dissolved oxygen were measured three times daily, whereas pH was measured once weekly (Table 3). All evaluated parameters remained within ranges considered suitable for Nile tilapia farming (²¹).

Table 3
Water quality parameters of experimental tanks containing juvenile Nile tilapia fed diets with increasing levels of ora-pro-nóbis leaf meal.

Time	Variable	Inclusion levels (%)					cv%	p-value
		0	3.5	7	10.5	14
Morning	Temperature	25.65	26.45	24.20	24.86	26.25	7.15	0.40
	Oxygen (mg/L)	7.12	7.45	6.86	7.39	6.90	12.93	0.84
	pH	7.34	7.81	7.25	8.00	7.75	6.81	0.25
Afternoon	Temperature	28.47	26.14	27.06	27.60	26.90	5.16	0.24
	Oxygen (mg/L)	6.64	6.87	7.05	7.10	7.26	6.05	0.33
	pH	7.76	8.87	8.13	7.78	7.09	10.67	0.10

2.5 Statistical analysis

All statistical analyses were performed using R software (²²). Growth performance and body chemical composition data were analyzed using analysis of variance (one-way ANOVA). Duncan’s test was applied for mean comparisons when significant differences were detected (p ≤ 0.05).

3. Results

Table 4 presents average initial weight (AIW), productive performance parameters, and survival values of juvenile Nile tilapia fed increasing levels of OLM. Average AFW, AWG, and DWG values of fish fed diets containing 7 %, 10.5 %, and 14 % OLM were higher than those observed for other treatments (p ≤ 0.05). Dietary OLM inclusion significantly affected fillet weight (p ≤ 0.05), with the highest value observed at maximum OLM inclusion level, whereas the lowest values were recorded for control diet and 3.5 % OLM inclusion level. Other treatments presented intermediate values. Dietary OLM inclusion positively affected (p ≤ 0.05) daily feed intake of Nile tilapia, which was higher in all OLM treatments. Regarding FCR and PER, diets containing OLM resulted in poorer values (p ≤ 0.05), whereas no dietary OLM effect was observed on SURV.

Table 4
Productive performance and survival parameters of juvenile Nile tilapia fed diets with increasing levels of ora-pro-nóbis leaf meal.

Variable	Inclusion levels (%)					CV%	p-value
	0	3.5	7	10.5	14
AIW	14.06	14.23	14.07	14.25	13.83	3.34	0.73
AFW	77.88b	78.78b	84.39a	85.61a	86.18a	3.27	0.00
AFLW	23.68b	24.11b	25.84ab	25.97ab	27.97a	6.35	0.01
AWG	63.83b	64.54b	70.32a	71.37a	72.35a	4.02	0.00
DWG	1.10b	1.11b	1.21a	1.23a	1.24a	4.02	0.00
DFI	1.47b	1.76a	1.79a	1.82a	1.83a	2.99	0.00
FC	1.40b	1.69a	1.55a	1.58a	1.60a	5.37	0.00
PER	2.40b	1.97a	2.12a	2.07a	2.08a	5.28	0.00
SURV (%)	100.00	96.25	98.75	98.75	100.00	1.96	0.09

Table 5 presents body yields and somatic indices of juvenile Nile tilapia fed increasing levels of OLM. CYH, FY, HSI, and VSI were not affected by dietary OLM inclusion, whereas VFI was significantly affected (p ≤ 0.05), with best values observed from 7 % OLM inclusion onward.

Table 5
Body yields and somatic indices of juvenile Nile tilapia fed increasing levels of ora-pro-nóbis leaf meal.

Variable	Inclusion levels (%)					CV%	p-value
	0	3.5	7	10.5	14
CYH	84.90	87.23	86.41	86.53	85.04	2.12	0.34
FY	31.46	30.85	30.62	31.65	32.51	4.39	0.37
HSI	2.16	1.54	1.76	1.52	1.64	21.33	0.15
VSI	15.10	12.77	13.59	13.47	15.22	12.77	0.27
VFI	2.65a	1.98a	1.19b	1.18b	0.97b	30.26	0.00

Table 6 shows the approximate chemical composition of fillets, carcasses, and whole fish of juvenile Nile tilapia fed increasing levels of OLM. Moisture (MOI), crude protein (CP), ether extract (EE), and mineral matter (MM) contents of fillets were not affected by dietary OLM inclusion. Regarding carcasses, MOI, CP, and MM were not affected by dietary OLM, whereas EE was significantly affected (p ≤ 0.05), with lower values observed at highest OLM inclusion levels (10.5 % and 14 %). In whole fish, dietary OLM inclusion significantly affected CP content (p ≤ 0.05), with highest values observed from 7 % OLM inclusion onward. In contrast, MOI, EE, and MM contents of whole fish were not affected by dietary OLM inclusion.

Table 6
Chemical composition of fillets, carcasses, and whole fish of juvenile Nile tilapia fed increasing levels of ora-pro-nóbis leaf meal.

	Nutrients1	Inclusion levels (%)					CV%	p-value
		0	3.5	7	10.5	14
Fillet	MO	75.45	75.58	75.40	75.56	76.36	1.27	0.62
	CP	20.30	20.22	20.35	20.15	19.54	3.89	0.60
	EE	2.20	1.99	2.30	1.99	1.96	19.36	0.69
	MM	5.16	5.04	5.00	5.17	5.20	3.66	0.49
Carcass	MO	71.00	71.10	70.89	71.67	71.43	0.75	0.27
	CP	29.00	28.90	29.11	28.33	28.57	1.87	0.27
	EE	7.74a	7.61ab	7.61ab	7.08c	7.04c	4.68	0.03
	MM	11.78	11.89	11.81	12.45	12.27	4.42	0.32
Fish	MO	69.18	69.43	69.08	70.23	70.71	1.52	0.18
	CP	16.48b	15.94b	16.76a	16.91a	17.24a	2.90	0.02
	EE	9.28	9.16	8.61	8.54	8.16	9.48	0.34
	MM	11.13	11.22	10.92	10.67	10.66	6.61	0.72

• ¹
  MO - moisture; CP - crude protein; EE - ether extract; MM - mineral matter.
• ²
  CV - coefficient of variation. Means followed by different letters within same row differ significantly according to Duncan’s test (P < 0.05).

4. Discussion

Dietary OLM inclusion in diets for juvenile Nile tilapia, particularly at inclusion levels of 7.0 %, 10.5 %, and 14.0 %, positively affected (p ≤ 0.05) important productive performance parameters, especially AFW, AFLW, AWG, DWG, and DFI. However, opposite effects were observed for FCR and PER, in which dietary OLM inclusion negatively affected these parameters (p ≤ 0.05).

Regarding AFW, AWG, and DWG, present findings differ from those reported by Matos et al. (¹⁹) and Dias et al. (²³), who observed no effects of OLM or mulberry leaf meal (MLM) on growth performance of Nile tilapia. Another important parameter was AFLW, which was significantly higher (p ≤ 0.05) at 14 % dietary OLM inclusion compared with control diet and 3.5 % treatment, although values did not differ from 7 % and 10.5 % inclusion levels. This response may be associated with higher DFI observed in present study, which was not reported in previous studies. Increased feed intake likely enhanced nutrient and energy availability for fish growth.

An increase in DFI was observed in fish fed diets containing OLM. Inclusion levels ranging from 3.5 % to 14 % significantly increased feed intake compared with control diet without OLM (p ≤ 0.05), suggesting a possible effect on feed palatability or attractiveness. Although this was not primary focus of present study, these findings indicate need for further investigations regarding influence of OLM on feed acceptance by Nile tilapia.

Higher DFI did not result in proportional weight gain, negatively affecting FCR (p ≤ 0.05). Although OLM-containing diets increased AWG by approximately 11 %, DFI increased more markedly (approximately 22 %), explaining reduced feed efficiency. In contrast, Matos et al. (19) reported improved FCR in adult Nile tilapia fed diets containing up to 10 % OLM, which authors associated with improvements in intestinal morphology and nutrient absorption. Absence of histological evaluations in present study suggests that less developed intestinal microvilli in juveniles may have limited dietary utilization efficiency.

Protein efficiency ratio, which reflects efficiency of converting consumed protein into body weight gain, was significantly impaired (p ≤ 0.05) at all dietary OLM inclusion levels. Highest mean value was observed in control treatment, with no significant differences among OLM inclusion levels. This finding suggests reduced protein utilization compensated by increased feed intake.

Visceral fat index (VFI) was significantly affected by dietary OLM inclusion (p ≤ 0.05), with fish fed 7.0%, 10.5 %, and 14 % OLM diets presenting lower visceral fat deposition. Visceral fat accumulation currently represents an important issue in commercial Nile tilapia farming, and observed reductions should be further investigated, as they may indicate reduced lipid deposition and/or improved nutrient and energy utilization, directly influencing VFI.

Changes in VFI were directly reflected in whole fish chemical composition, as an inversely proportional relationship between lipid and protein deposition is generally observed. In this context, reduced VFI values resulting from dietary OLM inclusion increased whole fish crude protein content (p ≤ 0.05). However, this effect was not observed in fillets and carcasses since visceral fat is not included in these portions.

Dietary OLM inclusion also produced significant effects (p ≤ 0.05) on carcass EE and whole fish CP. Lower carcass EE values were observed in fish fed diets containing 10.5 % and 14 % OLM, whereas fish fed control diet presented highest values. Diets containing 3.5 % and 7.0 % OLM resulted in intermediate values. Regarding whole fish CP, highest values were observed in fish fed diets containing 7.0 %, 10.5 %, and 14.0 % OLM. These findings reinforce possibility of improved nutrient and energy utilization at higher OLM inclusion levels.

Shiau et al. (²⁴) evaluated dietary fiber inclusion (carboxymethyl cellulose) in diets for tilapia hybrids (Oreochromis niloticus × Oreochromis aureus) and also reported reduced lipid deposition in fish fed diets containing more than 6 % fiber. Similarly, Meurer et al. (²⁵) evaluated effects of dietary fiber (cellulose) in diets for Nile tilapia fingerlings and suggested that reduced nutrient absorption may be associated with decreased digesta transit time caused by increasing dietary fiber levels.

Furthermore, fibers may exert prebiotic effects by modulating intestinal microbiota and stimulating production of short-chain fatty acids (SCFAs), such as butyrate, which promotes intestinal mucosal integrity and enterocyte regulation (²⁶). In addition, antioxidant compounds may reduce oxidative stress, contributing to a more balanced and functional intestinal environment (²⁷).

Differences between present findings and those reported by Matos et al. (¹⁹) are likely associated with physiological stage of fish. Juveniles possess shorter intestines relative to body size, which increase proportionally with growth (²⁸), resulting in reduced exposure time of digesta to digestive enzymes and enterocytes. In addition, juveniles exhibit distinct digestive enzyme activity and lower efficiency in utilizing fiber and complex carbohydrates compared with adult fish (²⁹,³⁰). Furthermore, juvenile fish possess a still-developing intestinal microbiota, whereas adults exhibit a more stable and adapted microbial community (³¹). Together, these factors directly influence nutrient digestion and absorption in Nile tilapia at different developmental stages.

Diets containing OLM produced more positive than negative effects. However, interpretation of these findings is limited by scarcity of studies evaluating OLM inclusion in diets for Nile tilapia (¹¹,¹⁹), as well as by influence of factors beyond fiber and macronutrient composition. Ora-pro-nóbis contains bioactive compounds with antioxidant potential, including caftaric acid and flavonoids derived from quercetin, kaempferol, and isorhamnetin. In addition, antimicrobial activity against Gram-positive and Gram-negative bacteria has also been reported, which may contribute to improvements in intestinal health and productive performance (³²). Therefore, further studies evaluating OLM inclusion in diets for Nile tilapia are needed.

5. Conclusion

Ora-pro-nóbis leaf meal can be included at levels ranging between 7 and 14 % in diets for juvenile Nile tilapia without compromising survival, while improving growth performance and body composition, besides supporting a sustainable feed formulation.

Generative AI use statement

The authors did not use generative artificial intelligence tools or technologies in the creation or editing of any part of this manuscript.

Data availability statement

Data will be provided upon request by the corresponding author.

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Edited by

• Editor: Rondineli P. Barbero

Publication Dates

• Publication in this collection
  20 July 2026
• Date of issue
  2026

History

• Received
  21 Sept 2025
• Accepted
  24 Apr 2026
• Published
  28 May 2026