Multiple feature affixation in Seenku plural formation

Nominal plural formation in Seenku involves two surface changes: fronting of the final vowel and raising of the final tone. Diachronically, the plural patterns likely derive from a suffix *- Open image in new window , which has been obscured through the loss of falling sonority diphthongs. By comparing plural formation to other morphophonological processes in Seenku, this paper argues that the plural suffix has been restructured as a featural suffix consisting of two features: the vocalic feature [+front], resulting in vowel fronting, and the tonal feature [+raised], resulting in tone raising. Given the atomic nature of the morphosyntactic feature plural, Seenku plural formation represents a strong case of multiple feature affixation, albeit a case that can be accounted for through Max constraints on feature values (Lombardi 2001, etc.) and Realize-Morpheme (van Oostendorp 2005; Trommer 2012) rather than the constraint Max-Flt argued for by Wolf (2007). A level-ordered approach retaining the vocalic suffix is also considered but is shown to suffer from a number of shortcomings, particularly with respect to tone and a challenging class of nasal stems. In short, this paper explores how the synchronic grammar copes with the vestiges of affixal morphology in a language that has undergone heavy reduction.     

Plural exponence in the Nez Perce DP: a DM analysis

This paper analyzes two patterns of number marking in the DP in Nez Perce (Sahaptian) within the framework of Distributed Morphology. The first involves under-realization of plural on nouns. Number has classically been understood as a feature inherent to nouns, rather than to adjectives that modify them. In Nez Perce, however, only a small set of nouns show number morphology, whereas number morphology is highly productive on adjectival modifiers. Adjectives in fact may realize the plural more than once per word—an instance of multiple exponence. I show that the puzzle of under-realization for nouns can be solved through conditioned allomorphy, providing new evidence for the presence of gender features on n (Lowenstamm 2007; Kramer 2014). The puzzle of multiple exponence (over-realization) for adjectives can be solved through Local Dislocation (Embick and Noyer 2001) combined with very late insertion of reduplicative content (Haugen 2008, 2011), demonstrating one way the DM architecture may produce multiple exponence without recourse to dedicated mechanisms.     

Verb-noun compounds in Italian from the 16th century onwards: an increasing exploitation of an available word-formation pattern

Verb-noun compounds are typically considered to be one of the most important innovations in the domain of Romance word-formation (cf. Bauer 2011, among many others). This morphological innovation can also be seen as part of a general tendency in Romance languages to prefer head-initial structures across different levels of the language system (cf., e.g., Ledgeway 2012: 225). However, the question of the productivity of VN compounds arises immediately as there is evidence that the pattern has become extremely productive only recently. This paper investigates the question of the productivity of the VN compounds starting out with a distinction between availability of a word-formation pattern and its profitability (cf., e.g., Bauer 2001). This article shows that the structural presence—availability—of VN compounding seems to be diachronically constant, but that the quantitative exploitation of the pattern—its profitability—turns out to be a recent phenomenon. Following recent research on compounding in Spanish (Moyna 2011) and French (Rosenberg 2007, 2008, 2011), this paper takes one concrete example of the Italian VN compounds and proceeds to show its diachronic evolution from the 16th to the 19th century on the basis of data drawn from diachronic corpora as well as from major historical dictionaries. It demonstrates that the “dramatic increase” alluded to by Bauer (2011: 543) is not to be taken as a sign of a new structural innovation, but rather as a fortuitous exploitation of a well-settled word-formation pattern.     

From possessive suffix to affective demonstrative suffix in Hungarian: a grammaticalization analysis

The non-possessive uses of possessive morphology in Uralic languages have been a topic of intense debate (Fraurud 2001; Nikolaeva 2003; Gerland 2014; Janda 2015; É. Kiss and Tánczos to appear). In this paper, I focus on a special use of the poss.3sg suffix in Hungarian constructions such as a hülyéje (the stupid-poss.3sg): lit. ‘its stupid’, meaning ‘that total idiot’. My main claim is that this suffix is an affective demonstrative suffix (Lakoff 1974; Liberman 2008; Potts and Schwarz 2010), and that it has developed as a result of grammaticalization from a full-fledged possessive construction of the form a világ hülyéje (the world stupid-poss.3sg): lit. ‘the world’s stupid’, meaning: ‘the biggest idiot in the world’. I will show that this gradual process can be reconstructed fairly accurately using historical and contemporary corpora. I also claim that this grammaticalization pathway is very natural as it is based on a set-element relationship which is often expressed by possessive constructions cross-linguistically. I also identify two parameters which facilitate this grammaticalization process: the availability of (silent) pro possessors and the lack of gender agreement on the possessive suffix. Since Uralic languages in general have these parameters, I will argue that this grammaticalization pathway should at least be considered as one of the possible sources of the demonstrative (and definiteness marking) uses of poss.3sg suffixes in Uralic languages. Finally, my results are also an important contribution to the debate on whether demonstratives can be derived from other functional elements through grammaticalization (Plank 1979; Traugott 1982; Himmelmann 1997).     

A recurrent neural model for proto-object based contour integration and figure-ground segregation

Visual processing of objects makes use of both feedforward and feedback streams of information. However, the nature of feedback signals is largely unknown, as is the identity of the neuronal populations in lower visual areas that receive them. Here, we develop a recurrent neural model to address these questions in the context of contour integration and figure-ground segregation. A key feature of our model is the use of grouping neurons whose activity represents tentative objects (“proto-objects”) based on the integration of local feature information. Grouping neurons receive input from an organized set of local feature neurons, and project modulatory feedback to those same neurons. Additionally, inhibition at both the local feature level and the object representation level biases the interpretation of the visual scene in agreement with principles from Gestalt psychology. Our model explains several sets of neurophysiological results (Zhou et al. Journal of Neuroscience, 20(17), 6594–6611 2000; Qiu et al. Nature Neuroscience, 10(11), 1492–1499 2007; Chen et al. Neuron, 82(3), 682–694 2014), and makes testable predictions about the influence of neuronal feedback and attentional selection on neural responses across different visual areas. Our model also provides a framework for understanding how object-based attention is able to select both objects and the features associated with them.     

Lexical splits within periphrasis: mixed perfective auxiliation systems in Italo-Romance

This paper addresses the phenomenon of mixed paradigms, i.e. mixed perfective auxiliation systems, attested in a wide range of Italo-Romance varieties (cf. Loporcaro 2001, 2007a, 2014; Manzini and Savoia 2005, among others). In these varieties, two auxiliary verbs, esse and habere, alternate within one and the same (sub)paradigm, displaying various patterns which can range from morphosyntactically motivated to apparently unmotivated distributions (here termed ‘morphomic’). I propose that, in these varieties, auxiliary selection is no longer a syntactically driven phenomenon, but becomes morphologized. I draw on the notion of ‘lexical split’ (cf. Corbett 2013, 2015, 2016) and describe the attested splits induced by intraparadigmatic auxiliary alternation. Following Bonami (2015), I put forward a typology of such splits. It is shown that, apart from motivated distributions, some morphomic patterns can also be found. The typology becomes more complex insofar as patterns with free variation between both auxiliaries are taken into account, as well as patently morphomic patterns which also seem to display external syntactic relevance (cf. Corbett 2013: 174–176). The phenomena reviewed and discussed in this paper are of major interest because they demonstrate the existence of competing exponence strategies within periphrasis, thus enriching the notion of ‘possible lexeme’ (cf. Corbett 2015: 146).     

Plant individuality: a solution to the demographer’s dilemma

The problem of plant individuality is something which has vexed botanists throughout the ages, with fashion swinging back and forth from treating plants as communities of individuals (Darwin 1800; Braun and Stone 1853; Münch 1938) to treating them as organisms in their own right, and although the latter view has dominated mainstream thought most recently (Harper 1977; Cook 1985; Ariew and Lewontin 2004), a lively debate conducted mostly in Scandinavian journals proves that the issues are far from being resolved (Tuomi and Vuorisalo 1989b; Fagerström 1992; Pan and Price 2001). In this paper I settle the matter once and for all, by showing which elements of each side are correct.     

Segmental isotopic labeling of HIV-1 capsid protein assemblies for solid state NMR

Recent studies of noncrystalline HIV-1 capsid protein (CA) assemblies by our laboratory and by Polenova and coworkers (Protein Sci 19:716–730, 2010; J Mol Biol 426:1109–1127, 2014; J Biol Chem 291:13098–13112, 2016; J Am Chem Soc 138:8538–8546, 2016; J Am Chem Soc 138:12029–12032, 2016; J Am Chem Soc 134:6455–6466, 2012; J Am Chem Soc 132:1976–1987, 2010; J Am Chem Soc 135:17793–17803, 2013; Proc Natl Acad Sci USA 112:14617–14622, 2015; J Am Chem Soc 138:14066–14075, 2016) have established the capability of solid state nuclear magnetic resonance (NMR) measurements to provide site-specific structural and dynamical information that is not available from other types of measurements. Nonetheless, the relatively high molecular weight of HIV-1 CA leads to congestion of solid state NMR spectra of fully isotopically labeled assemblies that has been an impediment to further progress. Here we describe an efficient protocol for production of segmentally labeled HIV-1 CA samples in which either the N-terminal domain (NTD) or the C-terminal domain (CTD) is uniformly ¹⁵N,¹³C-labeled. Segmental labeling is achieved by trans-splicing, using the DnaE split intein. Comparisons of two-dimensional solid state NMR spectra of fully labeled and segmentally labeled tubular CA assemblies show substantial improvements in spectral resolution. The molecular structure of HIV-1 assemblies is not significantly perturbed by the single Ser-to-Cys substitution that we introduce between NTD and CTD segments, as required for trans-splicing.     

IFN-γ production in response to <Emphasis Type="Italic">in vitro</Emphasis> stimulation with collagen type II in rheumatoid arthritis is associated with HLA-DRB1<Superscript>*</Superscript>0401 and HLA-DQ8

Introduction

Despite much work over past decades, whether antigen-specific immune reactions occur in rheumatoid arthritis (RA) and to what extent such reactions are directed towards joint-specific autoantigens is still questionable. One strong indicator for antigenic involvement in RA is the fact that certain major histocompatibility complex (MHC) class II genotypes [human leucocyte antigen (HLA)-DR4 and HLA-DR1] predispose for the development of the disease [1]. In the present report, collagen type II (CII) was studied as a putative autoantigen on the basis of both clinical and experimental data that show an increased frequency of antibodies to CII in RA patients [2,3,4] and that show that CII can induce experimental arthritis [5].It is evident from the literature that RA peripheral blood mononuclear cells (PBMCs) respond poorly to antigenic stimulation [6,7,8], and in particular evidence for a partial tolerization to CII has been presented [9]. The strategy of the present work has accordingly been to reinvestigate T-cell reactivity to CII in RA patients, to relate it to the response to commonly used recall antigens and to analyze IFN-γ responses as an alternative to proliferative responses.     

Strong Comma-Free Codes in Genetic Information

Comma-free codes constitute a class of circular codes, which has been widely studied, in particular by Golomb et al. (Biologiske Meddelelser, Kongelige Danske Videnskabernes Selskab 23:1–34, 1958a, Can J Math 10:202–209, 1958b), Michel et al. (Comput Math Appl 55:989–996, 2008a, Theor Comput Sci 401:17–26, 2008b, Inf Comput 212:55–63, 2012), Michel and Pirillo (Int J Comb 2011:659567, 2011), and Fimmel and Strüngmann (J Theor Biol 389:206–213, 2016). Based on a recent approach using graph theory to study circular codes Fimmel et al. (Philos Trans R Soc 374:20150058, 2016), a new class of circular codes, called strong comma-free codes, is identified. These codes detect a frameshift during the translation process immediately after a reading window of at most two nucleotides. We describe several combinatorial properties of strong comma-free codes: enumeration, maximality, self-complementarity and \(CF^3\)-property (comma-free property in all the three possible frames). These combinatorial results also highlight some new properties of the genetic code and its evolution. Each amino acid in the standard genetic code is coded by at least one strong comma-free code of size 1. There are 9 amino acids \(S=\{Asn,Asp,Gln,Gly,Lys,Met,Phe,Pro,Trp\}\) among 20 such that for each amino acid from S, its synonymous trinucleotide set (excluding the necessary periodic trinucleotides \(\{AAA,CCC,GGG,TTT\}\)) is a strong comma-free code. The primeval comma-free RNY code of Eigen and Schuster (Naturwissenschaften 65:341–369, 1978) is a self-complementary \(CF^3\)-code of size 16. Furthermore, it is the union of two strong comma-free codes of size 8 which are complementary to each other.     

Chemical Analysis of a “Miller-Type” Complex Prebiotic Broth

In a famous experiment Stanley Miller showed that a large number of organic substances can emerge from sparking a mixture of methane, ammonia and hydrogen in the presence of water (Miller, Science 117:528–529, 1953). Among these substances Miller identified different amino acids, and he concluded that prebiotic events may well have produced many of Life’s molecular building blocks. There have been many variants of the original experiment since, including different gas mixtures (Miller, J Am Chem Soc 77:2351–2361, 1955; Oró Nature 197:862–867, 1963; Schlesinger and Miller, J Mol Evol 19:376–382, 1983; Miyakawa et al., Proc Natl Acad Sci 99:14,628–14,631, 2002). Recently some of Miller’s remaining original samples were analyzed with modern equipment (Johnson et al. Science 322:404–404, 2008; Parker et al. Proc Natl Acad Sci 108:5526–5531, 2011) and a total of 23 racemic amino acids were identified. To give an overview of the chemical variety of a possible prebiotic broth, here we analyze a “Miller type” experiment using state of the art mass spectrometry and NMR spectroscopy. We identify substances of a wide range of saturation, which can be hydrophilic, hydrophobic or amphiphilic in nature. Often the molecules contain heteroatoms, with amines and amides being prominent classes of molecule. In some samples we detect ethylene glycol based polymers. Their formation in water requires the presence of a catalyst. Contrary to expectations, we cannot identify any preferred reaction product. The capacity to spontaneously produce this extremely high degree of molecular variety in a very simple experiment is a remarkable feature of organic chemistry and possibly prerequisite for Life to emerge. It remains a future task to uncover how dedicated, organized chemical reaction pathways may have arisen from this degree of complexity.     

Mathematical Model of Bone Remodeling Captures the Antiresorptive and Anabolic Actions of Various Therapies

A better understanding of the molecular pathways regulating the bone remodeling process should help in the development of new antiresorptive regulators and anabolic regulators, that is, regulators of bone resorption and of bone formation. Understanding the mechanisms by which parathyroid hormone (PTH) influences bone formation and how it switches from anabolic to catabolic action is important for treating osteoporosis (Poole and Reeve in Curr Opin Pharmacol 5:612–617, 2005). In this paper we describe a mathematical model of bone remodeling that incorporates, extends, and integrates several models of particular aspects of this biochemical system (Cabal et al. in J Bone Miner Res 28(8):1830–1836, 2013; Lemaire et al. in J Theor Biol 229:293–309, 2004; Peterson and Riggs in Bone 46:49–63, 2010; Raposo et al. in J Clin Endocrinol Metab 87(9):4330–4340, 2002; Ross et al. in J Disc Cont Dyn Sys Series B 17(6):2185–2200, 2012). We plan to use this model as a bone homeostasis platform to develop anabolic and antiresorptive compounds. The model will allow us to test hypotheses about the dynamics of compounds and to test the potential benefits of combination therapies. At the core of the model is the idealized account of osteoclast and osteoblast signaling given by Lemaire et al. (J Theor Biol 229:293–309, 2004). We have relaxed some of their assumptions about the roles of osteoprotegerin, transforming growth factor \(\upbeta \), and receptor activator of nuclear factor \(\upkappa \)B ligand; we have devised more detailed models of the interactions of these species. We have incorporated a model of the effect of calcium sensing receptor antagonists on remodeling (Cabal et al. in J Bone Miner Res 28(8):1830–1836, 2013). We have also incorporated a basic model of the effects of vitamin D on calcium homeostasis. We have included a simple model of the mechanism proposed by Bellido et al. (2003), Ross et al. (J Disc Cont Dyn Sys Series B 17(6):2185–2200, 2012), of the influence of PTH on osteoblast apoptosis, a mechanism that accounts for the anabolic response to pulsatile PTH administration. Finally, we have devised a simple model of the administration and effects of bisphosphonates. The biomarkers in the model are procollagen type 1 amino-terminal propeptide and C-terminal telopeptide. Bone mineral density is the model’s principal endpoint.     

Use of CD40L immunoconjugates to overcome the defective immune response to vaccines for infections and cancer in the aged

Multiple investigators have reported thepresence of defects in the immune response of the elderly [Castle In: Clin Infect Dis 31:578, 2000; Ortqvist et al. In: Eur Respir J 30:414–422, 2007; Saurwein-Teissl et al. In: J Immunol 168:5893, 2002; Haynes et al. In: Proc Natl Acad Sci USA 100:15053–15058, 2003]. These defects reduce the magnitude of the immune response to infection and to vaccination. In individuals greater than 55 years of age, the probability of developing a fully protective neutralizing antibody response to the yearly multivalent particle inactivated influenza vaccine is less than 20% [Jefferson et al. In: Lancet 264:1165–1174, 2005; Goodwin et al. In: Vaccine 24:1159–1169, 2006; Jackson et al. In: Lancet 372:398–405, 2008; Simonsen and Taylor In: Lancet 7:658–666, 2007]. The defects in the aged immune system that are responsible for this limited response to vaccination in the older age groups include functional defects of the antigen presenting cells, functional defects in CD4 helper CD4 T cells and monocytes, and an altered microenvironment [Eaton et al. In: J Exp Med 200:1613–1622, 2004; Dong et al. In: J Gen Virol 84:1623–1628, 2003; Deng et al. In: Immunology 172:3437–3446, 2004; Cella et al. In: J Exp Med 184:747–752, 1996]. Starting at puberty, the involution of the thymus and the consequent reduction of the export of naïve T cells specific to neo-antigens leads to the reduction of the ratio of antigen naïve to memory cells as chronological age advances [Prelog In: AutoimmunRev 5:136–139, 2006; McElhaney et al. In: J Immunology 176:6333–6339, 2006]. Changes in glycosylation of T cells and target antigens acquired during the aging process and the antibodies to these new glycopeptides and glycoproteins may also contribute to a reduction in the functioning of the adaptive immune response [Ishii et al. In: J Clin Neurosci 14:110–115, 2007; Shirai et al. In: Clin Exp Immunol 12:455–464, 1972; Adkins and Riley In: Mech AgeingDev 103:147–164, 1998; Ben-Yehuda and Weksler In: Cancer Investigation 10:525–531, 1992]. One of the more interesting examples of the functional defects in the cells of the adaptive immune response is a reduced level of expression in the surface cytoadhesion and activation receptor molecules on CD4 helper T cells undergoing activation during vaccination. Upon infection or vaccination, CD40L is typically increased on the surface of CD4 helper T cells during activation, and this increased expression is absolutely essential to the CD40L promotion of expansion of antigen-specific B cells and CD 8 effector T cells in response to infection or vaccination [Singh et al. In: Protein Sci 7:1124–1135, 1998; Grewal and Flavell In: Immunol Res 16: 59–70, 1997; Kornbluth In: J Hematother Stem Cell Res 11:787–801, 2002; Garcia de Vinuesa et al. In: Eur J Immunol 29:3216–3224, 1999]. In aged human beings and mice, the reduced levels of expression of CD40 ligand (CD40L) in activated CD4 helper T cells is dramatically reduced [Eaton et al. In: J Exp Med 200:1613–1622, 2004; Dong et al. In: J Gen Virol 84:1623–1628, 2003]. To circumvent the reduction in CD40L expression and the subsequent reduction in immune response in the elderly, we have developed a chimeric vaccine comprised of the CD40L linked to the target antigen, in a replication incompetent adenoviral vector and in booster protein. This review will discuss the implementation the potential use of this approach for the vaccination of the older populations for cancer and infection.     

Weak radicals,weak suppletion,and phonological indices in Semitic

Paster (2014) has argued that grammar does not distinguish between weak and strong suppletion. This claim is countered in this paper, in an analysis of allomorphic effects in several Semitic languages. What has been traditionally called “root” is broken down into three levels: the formless index (Harley 2014), the phonological index (Borer 2005, 2009, 2013) and its underlying representation. The distinction between the first two leads to a principled structural difference between weak and strong suppletion. Several test cases are presented that require a distinction between the latter two. Two general conclusions are drawn: (1) allomorphy at the different levels has different functions, and (2) in the Semitic root-and-pattern system, weakly suppletive allomorphy never targets the root as a whole; there is no root allomorphy in Semitic.     

Production of Estonian case-inflected nouns shows whole-word frequency and paradigmatic effects

Most psycholinguistic models of lexical processing assume that the comprehension and production of inflected forms is mediated by morphemic constituents. Several more recent studies, however, have challenged this assumption by providing empirical evidence that information about individual inflected forms and their paradigmatic relations is available in long-term memory (Baayen et al. 1997; Milin et al. 2009a, 2009b). Here, we investigate how whole-word frequency, inflectional paradigm size and morphological family size affect production latencies and articulation durations when subjects are asked to read aloud isolated Estonian case-inflected nouns. In Experiment 1, we observed that words with a larger morphological family elicited shorter speech onset latencies, and that forms with higher whole-word frequency had shorter acoustic durations. Experiment 2, for which we increased statistical power by using 2,800 words, revealed that higher whole-word frequency, inflectional paradigm size, and morphological family size reduced both speech onset times and acoustic durations. These results extend our knowledge of morphological processing in three ways. First, whole-word frequency effects of inflected forms in morphologically rich languages are not restricted to a small number of very high-frequency forms, contrary to previous claims (Niemi et al. 1994; Hankamer 1989; Yang 2016). Second, we replicated the morphological family size effect in a new domain, the acoustic durations of inflected forms. Third, we showed that a novel paradigmatic measure, inflectional paradigm size, predicts word naming latencies and acoustic durations. These results fit well with Word-and-Paradigm morphology (Blevins 2016) and argue against strictly (de)compositional models of lexical processing.     

Aspect and modality in the interpretation of deverbal -<Emphasis Type="Italic">er</Emphasis> nominals in English

This article presents a corpus study of over 16,000 tokens of -er nominalizations on 62 verbal bases that were extracted from the Corpus of Contemporary American English and the British National Corpus. We show that an individual -er nominal can often be given a range of modal and aspectual readings and that a number of factors influence the availability of different readings for -er nominals, including verb type, syntactic context (verb tenses, adverbs), and encyclopedic information. On the basis of these data, we argue, contra Cohen (2016), that the core meaning of the affix -er (as in writer, printer, etc.) cannot be that of a dynamic modal. We show that neither Cohen’s (2016) analysis nor syntactic analyses such as that of Alexiadou and Schäfer (2010) can account for the range of readings we find. We conclude by sketching one possible analysis in terms of the Lexical Semantic Framework of Lieber (2004, 2016) that postulates underspecified lexical representations of the -er nominals and resolution of underspecification in context.     

<Emphasis Type="Italic">Djombangia mannai</Emphasis> sp. n. (Cestoidea: Caryophyllidea: Lytocestidae) from a Siluroid Fish in West Bengal,India

The genus Djombangia Bovien, 1926 is a monotypic genus with only one valid species Djombangia penetrans Bovien, 1926 under it. In this paper Djombangia mannai sp.n. obtained from the intestine of a siluriform fresh water fish Clarias batrachus Linnaeus, 1758 from Diamond Harbour, South-24 Parganas, West Bengal, India is described and illustrated. The species is characterized by a small, globular scolex with spherical apical organ; body without transverse grooves; absence of neck; absence of seminal receptacle and 300–350 testes that differentiates it from the only known species under this genus.     

First-principle investigation on growth patterns and properties of cobalt-doped lithium nanoclusters

A systematic theoretical investigation on cobalt lithium clusters Li_nCo [1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12] was performed with a DFT approach. The location of global minima and structural evolution were carried out using the partical swarm optimization method. Li₆Co is the transition structure in going from low-coordinated structures to three-dimensional torispherical structures with a cobalt atom enclosed by lithium atoms. Maxima of ?₂ E and E _b for Li_nCo were found at n?=?3, 6, 8, 10, indicating that these clusters possess higher relative stability than their neighbors. In comparison with small clusters, n?=?1–6, the greater electron transfer from Li-2s to Co-3d within cage-like clusters Li_nCo (n?=?7–12) strengthens the bonding effect between Li_n and Co, which is reflected in the Wiberg bond index of Co and atomic binding energy analysis. AdNDP analysis verified the presence of both Lewis bonding elements (1c–2e objects) and delocalized bonding elements (6c–2e, 9c–2e and 10c–2e bonds). It is hoped that this theoretical work will provide favorable information to help understand the influence of dopant transition metal atoms on the properties of lithium-based materials.     

Competition between phytoplankton and bacteria: exclusion and coexistence

Resource-based competition between microorganisms species in continuous culture has been studied extensively both experimentally and theoretically, mostly for bacteria through Monod and Contois “constant yield” models, or for phytoplankton through the Droop “variable yield” models. For homogeneous populations of N bacterial species (Monod) or N phytoplanktonic species (Droop), with one limiting substrate and under constant controls, the theoretical studies indicated that competitive exclusion occurs: only one species wins the competition and displaces all the others (Armstrong and McGehee in Am Nat 115:151, 1980; Hsu and Hsu in SIAM J Appl Math 68:1600–1617, 2008). The winning species expected from theory is the one with the lowest “substrate subsistence concentration” \(s^{\star }\), such that its corresponding equilibrium growth rate is equal to the dilution rate \(D\). This theoretical result was validated experimentally with phytoplankton (Tilman and Sterner in Oecologia 61(2):197–200, 1984) and bacteria (Hansen and Hubell in Science 207(4438):1491–1493, 1980), and observed in a lake with microalgae (Tilman in Ecology 58(22):338–348, 1977). On the contrary for aggregating bacterial species described by a Contois model, theory predicts coexistence between several species (Grognard et al. in Discrete Contin Dyn Syst Ser B 8(1):73–93, 2007). In this paper we present a generalization of these results by studying a competition between three different types of microorganisms: planktonic (or free) bacteria (represented by a generalized Monod model), aggregating bacteria (represented by a Contois model) and free phytoplankton (represented by a Droop model). We prove that the outcome of the competition is a coexistence between several aggregating bacterial species with a free species of bacteria or phytoplankton, all the other free species being washed out. This demonstration is based mainly on the study of the substrate concentration’s evolution caused by competition; it converges towards the lowest subsistence concentration \(s^{\star }\), leading to three different types of competition outcomes: (1) the best free bacteria or phytoplankton competitor excludes all other species; (2) only some aggregating bacterial species coexist in the chemostat; (3) A coexistence between the single best free species, with one or several aggregating species.