A systematic review of mesenchymal stem cell secretome: Functional annotations, gene clusters and proteomics analyses for bone formation.

The regenerative capacity of mesenchymal stem cells (MSCs) is now attributed to their ability to release paracrine factors into the extracellular matrix that boost tissue regeneration, reduce inflammation and encourage healing. Understanding the MSC secretome is crucial for shifting the prototypic conventional cell-based therapies to cell-free regenerative treatments. This systematic review aimed to analyse the functional annotations of the secretome of human adult adipose tissue and bone marrow MSCs and unveil the gene clusters responsible for bone formation. Bioinformatics tools were used to identify the biological processes, molecular functions, hallmarks and KEGG pathways of adipose and bone marrow MSC secretome proteins. We found a substantial overlap in the functional annotations and protein compositions of both adipose and bone marrow MSC secretome indicating that MSC source may be noninfluencial with regards to tissue regeneration. Additionally, a novel network pharmacology-based analysis of the secreted proteins revealed that the commonly secreted proteins within a single source interact with multiple drugable targets of bone diseases and regulate various KEGG pathway. This study unravels the secretome profile of human adult adipose and bone marrow MSCs based on the current literature and provides valuable insights into the therapeutic use of the MSC secretome for cell-free therapies.

Coexistence of populations in a Leslie-Gower tritrophic model with Holling-type functional responses.

The dynamics of a Leslie-Gower type tritrophic model are analyzed. The model considers the interaction among three populations, a resource, a predator and a superpredator. It is assumed that the predator is generalist and its interaction with the resource is according to a general Holling-type functional response. Furthermore, it is assumed that the superpredator is specialist and its interaction with the predator follows a Holling type II functional response. The goal of this work is to show conditions that guarantee the coexistence of the three species. To do this, the existence of a stable equilibrium point or a stable limit cycle is demonstrated, which appears via a bifurcation. In addition, the analytical results are exemplified through numerical simulations.

Assessing potential for biological control of Japanese giant silkworm Caligula japonica using Anastatus gansuensis, a thelytokous parasitoid firstly reported in Eupelmidae.

BACKGROUND: Caligula japonica, commonly known as Japanese giant silkworm (JGS), is a serious defoliating pest of fruit and forest trees in East Asia. To develop eco-friendly and cost-effective control methods for this pest, we evaluated the potential for biological control of JGS using its egg parasitoid Anastatus gansuensis reared on the Chinese oak silkworm (COS) Antheraea pernyi. We compared the reproductive traits and population increase potential of the parasitoid on JGS and COS eggs, as well as its functional response to host egg densities and mutual interference at different parasitoid densities. RESULTS: Anastatus gansuensis was confirmed to be strictly synovigenic, with most eggs maturing post-emergence, and produced <1% male offspring on both host eggs. Although A. gansuensis females reared from COS eggs had longer longevity and oviposition period, and higher fecundity and net reproductive rate compared to those reared from JGS eggs, the parasitoid had a higher intrinsic rate of increase on JGS than COS eggs. The parasitoid exhibited a type II functional response to increasing host densities, with mutual interference among foraging female wasps occurred at higher parasitoid densities. CONCLUSION: Our results indicate a high potential for biological control of JGS using A. gansuensis. The parasitoid can be efficiently reared on COS eggs and used against JGS. It may be essential to provide food for emerging adult parasitoids allowing time for egg maturation prior to the rearing or augmentative release of the parasitoid. Some mutual interference at high parasitoid rearing densities likely reduces per capita parasitization efficiency of A. gansuensis. © 2024 Society of Chemical Industry.

Modeling the differential functional responses and selectivity of a marine copepod to nano/microplastics in mixture.

Nano- and microplastics (NMPs) pollution is widespread in the oceans, posing potential risks to marine species. This study examined the accumulation capacity and selectivity potentials of NMPs by a marine copepod Parvocalanus crassirostris under different food mixtures by modeling the combined biokinetic and functional response. We investigated two sizes of NMPs (200 nm and 5 µm) across a concentration gradient (0 - 5000 µg/L) and varying diatom abundances (0, 104, 105 cells/mL). Fluorescence imaging and quantification revealed that P. crassirostris actively ingested NMPs at low concentration. Accumulation increased with NMPs concentration but eventually saturated due to gut capacity limits, following a Holling type II functional response (i.e., hyperbolic curve). Our novel functional response model estimated the key parameters and demonstrated that the maximum accumulation reached 5.3 % of dry weight with averaged half-saturation constants of 229 µg/L. The size of NMPs did not significantly affect the total accumulation or satiety levels. The presence of diatoms influenced the feeding selectivity and decreased the microplastic accumulation by 73 % at 105 cells/mL, while facilitating nanoplastic accumulation by 81 % at 104 cells/mL. This study enhanced our understanding of NMPs bioavailability and environmental fate in marine ecosystems.

Short-term temperature changes affected the predation ability of Orius similis on Bemisia tabaci nymphs.

Bemisia tabaci (Gennadius), a major pest that can adversely affect economies and agriculture globally, is particularly sensitive to climate change-induced temperature fluctuations, which can intensify its outbreaks. Orius similis Zheng, a primary natural predator of B. tabaci, also experiences temperature-related effects that influence its biocontrol efficacy. Thus, understanding the response of O. similis to temperature changes is pivotal for optimizing its biocontrol potential. Herein, our investigations showed that the functional response of O. similis to both high- and low-instar nymphs of B. tabaci adheres to the type II model at temperatures of 19, 22, 25, 28, and 31 °C. At 28 °C, O. similis exhibits the highest instantaneous attack rate (high-instar: 1.1580, low-instar: 1.2112), and the shortest handling time per prey (high-instar: 0.0218, low-instar: 0.0191). The efficacy of O. similis in controlling B. tabaci nymphs follows the sequence: 28 °C > 25 °C > 31 °C > 22 °C > 19 °C. Additionally, search efficiency inversely correlates with prey density. Simulations using the Hessell-Varley interference model indicate that increased density of O. similis under any temperature condition leads to reduced predation rates. Moreover, O. similis shows a predation preference for low-instar nymphs of B. tabaci, with higher predation level observed at the same temperature. In conclusion, for effective control of B. tabaci in field releases, O. similis should be optimally released at temperatures between 25 and 28 °C to preferably target the egg or early nymph stages of B. tabaci and determining the appropriate number of O. similis is important to minimize interference among individuals and enhance biocontrol efficacy.

Functional response metrics explain and predict high but differing ecological impacts of juvenile and adult lionfish.

Recent accumulation of evidence across taxa indicates that the ecological impacts of invasive alien species are predictable from their functional response (FR; e.g. the maximum feeding rate) and functional response ratio (FRR; the FR attack rate divided by handling time). Here, we experimentally derive these metrics to predict the ecological impacts of both juvenile and adult lionfish (Pterois volitans), one of the world's most damaging invaders, across representative and likely future prey types. Potentially prey-population destabilizing Type II FRs were exhibited by both life stages of lionfish towards four prey species: Artemia salina, Gammarus oceanicus, Palaemonetes varians and Nephrops norvegicus. FR magnitudes revealed ontogenetic shifts in lionfish impacts where juvenile lionfish displayed similar if not higher consumption rates than adult lionfish towards prey, apart from N. norvegicus, where adult consumption rate was considerably higher. Additionally, lionfish FRR values were very substantially higher than mean FRR values across known damaging invasive taxa. Thus, both life stages of lionfish are predicted to contribute to differing but high ecological impacts across prey communities, including commercially important species. With lionfish invasion ranges currently expanding across multiple regions globally, efforts to reduce lionfish numbers and population size structure, with provision of prey refugia through habitat complexity, might curtail their impacts. Nevertheless, the present study indicates that management programmes to support early detection and complete eradication of lionfish individuals when discovered in new regions are advised.

Behavioural response to boat noise weakens the strength of a trophic link in coral reefs.

In oceans, the noise generated by human activities has reached phenomenal proportions, with considerable harmful effects on marine life. Measuring this impact to achieve a sustainable balance for highly vulnerable marine ecosystems, such as coral reefs, is a critical environmental policy objective. Here, we demonstrate that anthropogenic noise alters the interactions of a coral reef fish with its environment and how this behavioural response to noise impairs foraging. In situ observations on the Moorea reef revealed that the damselfish Dascyllus emamo reacts to boat passage by moving closer to its coral bommie, considerably reducing the volume of water available to search for prey. Using boat noise playback experiments in microcosms, we studied D. emamo's behaviour and modeled its functional response (FR), which is the relationship between resource use and resource density, when feeding on juvenile shrimps. Similar to field observations, noise reduced D. emamo's spatial occupancy, accompanied by a lower FR, indicating a reduction in predation independent of prey density. Overall, noise-induced behavioural changes are likely to influence predator-prey interaction dynamics and ultimately the fitness of both protagonists. While there is an urgent need to assess the effect of anthropogenic noise on coral reefs, the ecological framework of the FR approach combined with behavioural metrics provides an essential tool for evaluating the cascading effects of noise on nested ecological interactions at the community level.

Resilience mechanisms of Trichopria drosophilae (Hymenoptera: Diapriinae) under global extreme cooling: insights into parasitic response and physiological adaptation.

Global climate warming and frequent extreme low-temperature events have made it essential to investigate the impact of low temperatures on parasitic wasps to protect and strengthen farmland biodiversity, which in turn enhances the biological control potential of natural enemies such as parasitic wasps. We systematically examined how low-temperature stress affects the parasitic functional response of Trichopria drosophilae to Drosophila suzukii (Diptera: Drosophilidae) pupae. Our findings indicate that the parasitic behavior of T. drosophilae towards D. suzukii pupae aligns with the Holling II functional response model following exposure to different temperatures. Within the temperature range of 8 °C to -8 °C, lower temperatures correlated decreased instantaneous attack rate of T. drosophilae and an increase in processing time. The search constant Q initially increased and then decreased with declining temperatures. Short-term low-temperature stress negatively impacted the parasitic and searching abilities of T. drosophilae but did not alter its parasitic functional response model. Notably, short-term low-temperature stress had minimal effects on the water content, protein content, and total sugar content of male and female T. drosophilae adults. However, as temperatures decreased, the activities of key enzymes, including GAPDH, SOD, T-AOC, and malondialdehyde (MDA), exhibited an initial increase followed by a decrease. Conversely, the activities of LDH and HOAD decreased, while the activities of CAT and POD increased. Further study on the effect of short-term low temperature on T. drosophilae can provide a research basis for the large-scale production and low-temperature refrigeration technology of T. drosophilae, and provide a scientific basis for its efficient use in the field.

Carnivore space use behaviors reveal variation in responses to human land modification.

BACKGROUND: Spatial behavior, including home-ranging behaviors, habitat selection, and movement, can be extremely informative in estimating how animals respond to landscape heterogeneity. Responses in these spatial behaviors to features such as human land modification and resources can highlight a species' spatial strategy to maximize fitness and minimize mortality. These strategies can vary on spatial, temporal, and individual scales, and the combination of behaviors on these scales can lead to very different strategies among species. METHODS: Harnessing the variation present at these scales, we characterized how species may respond to stimuli in their environments ranging from broad- to fine-scale spatial responses to human modification in their environment. Using 15 bobcat-years and 31 coyote-years of GPS data from individuals inhabiting a landscape encompassing a range of human land modification, we evaluated the complexity of both species' responses to human modification on the landscape through their home range size, habitat selection, and functional response behaviors, accounting for annual, seasonal, and diel variation. RESULTS: Bobcats and coyotes used different strategies in response to human modification in their home ranges, with bobcats broadly expanding their home range with increases in human modification and displaying temporal consistency in functional response in habitat selection across both season and time of day. Meanwhile, coyotes did not expand their home ranges with increased human modification, but instead demonstrated fine-scale responses to human modification with habitat selection strategies that sometimes varied by time of day and season, paired with functional responses in selection behaviors. CONCLUSIONS: These differences in response to habitat, resources, and human modification between the two species highlighted the variation in spatial behaviors animals can use to exist in anthropogenic environments. Categorizing animal spatial behavior based on these spatiotemporal responses and individual variation can help in predicting how a species will respond to future change based on their current spatial behavior.

Effect of Sex, Age and Temperature on the Functional Response of Macrolophus pygmaeus Ramber and Nesidiocoris tenuis Reuter (Heteroptera: Miridae) on Eggs of Tuta absoluta.

The predatory mirids Macrolophus pygmaeus Ramber (Heteroptera: Miridae) and Nesidiocoris tenuis Reuter (Heteroptera: Miridae) are used for the biological control of Tuta absoluta Meyrick (Lepidoptera: Gelechiidae) and other pests in tomato greenhouses. The functional response of 1-day-old (young) and 10-day-old (old) adult females and males of M. pygmaeus and N. tenuis on eggs of T. absoluta was determined on tomato at two temperatures (20 °C and 25 °C) and LD 16:8. Females of M. pygmaeus exhibited higher predation efficiency than males at both tested temperatures. Young M. pygmaeus females had a higher efficiency than old ones, whereas males had a low efficiency irrespective of age. The predation efficiency of N. tenuis was high (but lower than M. pygmaeus) in both young females and males, although old females had a higher efficiency than the respective males. Our results show that the two predatory species have different functional response characteristics to their prey depending mainly on sex and age, which may affect their role as biological control agents.

Importance of Host Feeding in the Biological Control of Insect Pests: Case Study of Egg Parasitoid Species (Hymenoptera: Chalcidoidea: Trichogrammatidae).

Over recent decades, intraguild predation (IGP) has attracted special attention, both from the theoretical and practical standpoints. The present paper addresses the interference competition between two Trichogramma species (egg parasitoids)-on the one hand, the extrinsic interactions (i.e., the indirect competition between female T. achaeae and T. brassicae), and on the other, the intrinsic interactions between the larvae of both species. Furthermore, T. achaeae is a better competitor than T. brassicae due to a dual mechanism-the former acts as a facultative hyperparasitoid of the latter, exclusively considering parasitism relationships as well as presenting predation activity by host feeding, which gives preference to eggs previously parasitized by T. brassicae over non-parasitized eggs. Both mechanisms are dependent on the prey density, which is demonstrated by a change in the functional response (i.e., the relationship between the numbers of prey attacked at different prey densities) of T. achaeae adult female-it changes from type II (i.e., initial phase in which the number of attacked targets increases hyperbolically and then reaches an asymptote, reflecting the handling capacity of the predator), in the absence of competition (an instantaneous search rate of a' = 9.996 ± 4.973 days-1 and a handling time of Th = 0.018 ± 0.001 days), to type I (i.e., linear increase in parasitism rate as host densities rise, until reaching a maximum parasitism rate, and an instantaneous search rate of a' = 0.879 ± 0.072 days-1 and a handling time of Th ≈ 0) when interference competition is present. These results show that there is a greater mortality potential of this species, T. achaeae, in conditions of competition with other species, T. brassicae in this case. Based on this, their implications in relation to the biological control of pests by parasitoid species are discussed.

Food web context modifies predator foraging and weakens trophic interaction strength.

Trophic interaction modifications (TIM) are widespread in natural systems and occur when a third species indirectly alters the strength of a trophic interaction. Past studies have focused on documenting the existence and magnitude of TIMs; however, the underlying processes and long-term consequences remain elusive. To address this gap, we experimentally quantified the density-dependent effect of a third species on a predator's functional response. We conducted short-term experiments with ciliate communities composed of a predator, prey and non-consumable 'modifier' species. In both communities, increasing modifier density weakened the trophic interaction strength, due to a negative effect on the predator's space clearance rate. Simulated long-term dynamics indicate quantitative differences between models that account for TIMs or include only pairwise interactions. Our study demonstrates that TIMs are important to understand and predict community dynamics and highlights the need to move beyond focal species pairs to understand the consequences of species interactions in communities.

The environment to the rescue: can physics help predict predator-prey interactions?

Understanding the factors that determine the occurrence and strength of ecological interactions under specific abiotic and biotic conditions is fundamental since many aspects of ecological community stability and ecosystem functioning depend on patterns of interactions among species. Current approaches to mapping food webs are mostly based on traits, expert knowledge, experiments, and/or statistical inference. However, they do not offer clear mechanisms explaining how trophic interactions are affected by the interplay between organism characteristics and aspects of the physical environment, such as temperature, light intensity or viscosity. Hence, they cannot yet predict accurately how local food webs will respond to anthropogenic pressures, notably to climate change and species invasions. Herein, we propose a framework that synthesises recent developments in food-web theory, integrating body size and metabolism with the physical properties of ecosystems. We advocate for combination of the movement paradigm with a modular definition of the predation sequence, because movement is central to predator-prey interactions, and a generic, modular model is needed to describe all the possible variation in predator-prey interactions. Pending sufficient empirical and theoretical knowledge, our framework will help predict the food-web impacts of well-studied physical factors, such as temperature and oxygen availability, as well as less commonly considered variables such as wind, turbidity or electrical conductivity. An improved predictive capability will facilitate a better understanding of ecosystem responses to a changing world.

Effects of field releases of Neoseiulus barkeri on Megalurothrips usitatus abundance and arthropod diversity.

Megalurothrips usitatus (Bagnall) (Thysanoptera: Thripidae) is an important pest in Vigna unguiculata (L.) Walp. Neoseiulus barkeri (Hughes) (Acari: Phytoseiidae) is widely used for control of pest mites and insects worldwide. We evaluated its effect on M. usitatus when predators (N. barkeri) or insecticides (Spinetoram) were applied in the fields. Neoseiulus barkeri Hughes consumed 80% of M. usitatus prey offered within 6 h, and predation showed Type III functional response with prey density. The maximum consumption of N. barkeri was 27.29 ± 1.02 individuals per d per arena (1.5 cm diameter), while the optimal prey density for the predatory mite was 10.35 ± 0.68 individuals per d per arena (1.5 cm diameter). The developmental duration of N. barkeri fed with M. usitatus was significantly shorter than those fed with the dried fruit mite, Carpoglyphus lactis (L.) (Acari: Astigmata). In field trials, the efficiency of N. barkeri against M. usitatus was not significantly different from that of applications of the insecticide spinetoram. Biodiversity of other insects in treated fields was assessed, and there were 21 insect species in garden plots treated with N. barkeri releases. The total abundance (N), Shannon's diversity index (H), Pielou's evenness index (J) and Simpson's diversity index (D) of the garden plots treated with predatory mites were all significantly higher than that in the garden plots treated with spinetoram, where we found no species of predators or parasitoids and 7 herbivores. Our results show that N. barkeri is a potential means to control M. usitatus while preserving arthropod diversity at the level of treated gardens.

Implications of Temperature and Prey Density on Predatory Mite Amblyseius swirskii (Acari: Phytoseiidae) Functional Responses.

Amblyseius swirskii are predaceous mites that feed on phytophagous mites, pollens, and plant exudates and are known as one of the most potent biological pest management agents. Tetranychus urticae is a global mite that is difficult to manage because of its high population growth rates, necessitating alternative management measures like biological control. Regarding the functional response, the effects of temperature and prey density are some of the essential behaviors of natural enemies. This study investigates the effect of varying temperatures and prey densities on A. swirskii, a biological control agent for T. urticae. The present results demonstrated the change in the functional response estimates when A. swirskii was reared at various temperatures and different prey densities. The results of the estimates regarding the searching efficiency (a') showed the highest value (a' = 0.919) at 26 °C and the lowest value (a' = 0.751) at 14 °C. The handling time per prey item (Th) for the predatory mites changed with the temperature and prey density, showing the shortest handling time at 26 °C (Th = 0.005) and the highest value at 14 °C (Th = 0.015). The functional response curves matched the type II functional response model, demonstrating the inverse dependence of temperatures and prey density with a positive quadratic coefficient. The predation curves for A. swirskii showed a significant difference between the mean numbers of T. urticae consumed at various prey densities and temperatures, illustrating a relationship between A. swirskii and T. urticae. Therefore, the results of this research may be utilized to forecast the behavior of A. swirskii and its usefulness in controlling T. urticae populations.

A taste of space: Remote animal observations and discrete-choice models provide new insights into foraging and density dynamics for a large subarctic herbivore.

Competition for resources and space can drive forage selection of large herbivores from the bite through the landscape scale. Animal behaviour and foraging patterns are also influenced by abiotic and biotic factors. Fine-scale mechanisms of density-dependent foraging at the bite scale are likely consistent with density-dependent behavioural patterns observed at broader scales, but few studies have directly tested this assertion. Here, we tested if space use intensity, a proxy of spatiotemporal density, affects foraging mechanisms at fine spatial scales similarly to density-dependent effects observed at broader scales in caribou. We specifically assessed how behavioural choices are affected by space use intensity and environmental processes using behavioural state and forage selection data from caribou (Rangifer tarandus granti) observed from GPS video-camera collars using a multivariate discrete-choice modelling framework. We found that the probability of eating shrubs increased with increasing caribou space use intensity and cover of Salix spp. shrubs, whereas the probability of eating lichen decreased. Insects also affected fine-scale foraging behaviour by reducing the overall probability of eating. Strong eastward winds mitigated negative effects of insects and resulted in higher probabilities of eating lichen. At last, caribou exhibited foraging functional responses wherein their probability of selecting each food type increased as the availability (% cover) of that food increased. Space use intensity signals of fine-scale foraging were consistent with density-dependent responses observed at larger scales and with recent evidence suggesting declining reproductive rates in the same caribou population. Our results highlight potential risks of overgrazing on sensitive forage species such as lichen. Remote investigation of the functional responses of foraging behaviours provides exciting future applications where spatial models can identify high-quality habitats for conservation.

Field and laboratory microplastics uptake by a freshwater shrimp.

Microplastics are widespread pollutants, but few studies have linked field prevalence in organisms to laboratory uptakes. Aquatic filter feeders may be particularly susceptible to microplastic uptake, with the potential for trophic transfer to higher levels, including humans. Here, we surveyed microplastics from a model freshwater shrimp, common caraidina (Caridina nilotica) inhabiting the Crocodile River in South Africa to better understand microplastic uptake rates per individual. We then use functional response analysis (feeding rate as a function of resource density) to quantify uptake rates by shrimps in the laboratory. We found that microplastics were widespread in C. nilotica, with no significant differences in microplastic abundances among sampled sites under varying land uses, with an average abundance of 6.2 particles per individual. The vast majority of microplastics found was fibres (86.1%). Shrimp microplastic accumulation patterns were slightly higher in the laboratory than the field, where shrimp exhibited a hyperbolic Type II functional response model under varying exposure concentrations. Maximum feeding rates of 20 particles were found over a 6 h feeding period, and uptake evidenced at even the lowest laboratory concentrations (~10 particles per mL). These results highlight that microplastic uptake is widespread in field populations and partly density dependent, with field concentrations corroborating uptake rates recorded in the laboratory. Further research is required to elucidate trophic transfer from these taxa and to understand potential physiological impacts.

Does mutual interference stabilize prey-predator model with Bazykin-Crowley-Martin trophic function?

We investigated a system of ordinary differential equations that describes the dynamics of prey and predator populations, taking into account the Allee effect affecting the reproduction of the predator population, and mutual interference amongst predators, which is modeled with the Bazykin-Crowley-Martin (BCM) trophic function. Bifurcation analysis revealed a rich spectrum of bifurcations occurring in the system. In particular, analytical conditions for the saddle-node, Hopf, cusp, and Bogdanov-Takens bifurcations were derived for the model parameters, quantifying the strength of the predator interference, the Allee effect, and the predation efficiency. Numerical simulations verify and illustrate the analytical findings. The main purpose of the study was to test whether the mutual interference in the model with BCM trophic function provides a stabilizing or destabilizing effect on the system dynamics. The obtained results suggest that the model demonstrates qualitatively the same pattern concerning varying the interference strength as other predator-dependent models: both low and very high interference levels increase the risk of predator extinction, while moderate interference has a favorable effect on the stability and resilience of the prey-predator system.

Foraging rates from metabarcoding: Predators have reduced functional responses in wild, diverse prey communities.

Functional responses describe foraging rates across prey densities and underlie many fundamental ecological processes. Most functional response knowledge comes from simplified lab experiments, but we do not know whether these experiments accurately represent foraging in nature. In addition, the difficulty of conducting multispecies functional response experiments means that it is unclear whether interaction strengths are weakened in the presence of multiple prey types. We developed a novel method to estimate wild predators' foraging rates from metabarcoding data and use this method to present functional responses for wild wolf spiders foraging on 27 prey families. These field functional responses were considerably reduced compared to lab functional responses. We further find that foraging is sometimes increased in the presence of other prey types, contrary to expectations. Our novel method for estimating field foraging rates will allow researchers to determine functional responses for wild predators and address long-standing questions about foraging in nature.

Effects of whaling and krill fishing on the whale-krill predation dynamics: bifurcations in a harvested predator-prey model with Holling type I functional response.

In the Antarctic, the whale population had been reduced dramatically due to the unregulated whaling. It was expected that Antarctic krill, the main prey of whales, would grow significantly as a consequence and exploratory krill fishing was practiced in some areas. However, it was found that there has been a substantial decline in abundance of krill since the end of whaling, which is the phenomenon of krill paradox. In this paper, to study the krill-whale interaction we revisit a harvested predator-prey model with Holling I functional response. We find that the model admits at most two positive equilibria. When the two positive equilibria are located in the region { ( N , P ) | 0 ≤ N < 2 N c , P ≥ 0 } , the model exhibits degenerate Bogdanov-Takens bifurcation with codimension up to 3 and Hopf bifurcation with codimension up to 2 by rigorous bifurcation analysis. When the two positive equilibria are located in the region { ( N , P ) | N > 2 N c , P ≥ 0 } , the model has no complex bifurcation phenomenon. When there is one positive equilibrium on each side of N = 2 N c , the model undergoes Hopf bifurcation with codimension up to 2. Moreover, numerical simulation reveals that the model not only can exhibit the krill paradox phenomenon but also has three limit cycles, with the outmost one crosses the line N = 2 N c under some specific parameter conditions.