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- © 2015 American Society of Plant Biologists. All Rights Reserved.
Ethylene (ET) is an important hormone in plant responses to microbial pathogens and herbivorous insects, and in the interaction of plants with beneficial microbes and insects. Early ET signaling events during these biotic interactions involve activities of mitogen-activated protein kinases and ETHYLENE RESPONSE FACTOR transcription factors. Rather than being the principal regulator, ET often modulates defense signaling pathways, including those regulated by jasmonic acid and salicylic acid. Hormonal signal integrations with ET steer the defense signaling network to activate specific defenses that can have direct effects on attackers, or systemically prime distant plant parts for enhanced defense against future attack. ET also regulates volatile signals that attract carnivorous enemies of herbivores or warn neighboring plants. Conversely, ET signaling can also be exploited by attackers to hijack the defense signaling network to suppress effective defenses. In this review, we summarize recent findings on the significant role of ET in the plants' battle against their enemies.
Plants live in complex environments in which they are constantly exposed to a wide range of biotic interactors. Some of these interactors, such as mutualistic rhizosphere bacteria and fungi, are beneficial for the plant as they promote plant growth and protect the plant from attack by harmful interactors. Conversely, harmful interactors, such as herbivorous insects and microbial pathogens, reduce the fitness of the plant by retrieving energy-rich organic compounds without returning a net benefit to the plant (Pieterse et al., 2014b). To minimize the success of attack by other organisms, plants have evolved sophisticated defensive mechanisms that are either expressed constitutively or induced when an attacker is recognized. Active defense against biotrophic pathogens, which form a long-term relationship with living plant cells to derive nutrients, is mainly effectuated by programmed plant cell death (Glazebrook, 2005). Infection by necrotrophic pathogens, which first destroy host cells before feeding on the content, is usually repulsed by plant-produced antimicrobial compounds (Glazebrook, 2005). To defend themselves against insects, plants can activate both direct and indirect defenses. Direct defenses include plant traits that hinder the insect's growth rate, adult size, and/or survival probability, such as trichomes or toxic secondary metabolites (Howe and Jander, 2008). Indirect defenses include plant traits that enhance the probability of attracting natural enemies of herbivorous insects, such as volatile organic compounds or extrafloral nectar (Dicke, 2015; Heil, 2015).
Inducible defenses are initiated after perception of microbial infection or insect infestation, for which plants possess a suite of pattern recognition receptors (PRRs). These PRRs specifically recognize general nonself molecules from attacking organisms and self molecules from already attacked plant cells (Cook et al., 2015). By detecting highly conserved structures of entire classes of microbes, so-called microbe-associated molecular patterns (MAMPs), plants can recognize attack by a wide variety of potential pathogens. Similarly, general elicitors that are present in the saliva of insects function as herbivore-associated molecular patterns. Moreover, enzymatic degradation of plant material by attacking microbes or insects generates endogenous elicitors, so-called damage-associated molecular patterns (Ferrari et al., 2013; Savatin et al., 2014; Acevedo et al., 2015). All of these different molecular patterns are detected through a general detection system consisting of PRRs and coreceptors, leading to the activation of pattern-triggered immunity (PTI; Jones and Dangl, 2006). Biosynthesis of the gaseous hormone ethylene (ET) is among the suite of immediate PTI responses that, together with the production of reactive oxygen species and the activation of mitogen-activated protein kinase (MAPK) signaling cascades, regulate the production of downstream defensive proteins and metabolites (Boller and Felix, 2009; Wu and Baldwin, 2010). A second layer of more specific perception of microbes and insects is accomplished by plant resistance (R) proteins. Successful pathogens are able to suppress or evade PTI by the production of attacker-specific effectors (Pel and Pieterse, 2013). In turn, R proteins in the plant have evolved to specifically recognize these effectors, initiating effector-triggered immunity (ETI) or R gene-mediated resistance (Jones and Dangl, 2006; Cui et al., 2015). ETI is accompanied by rapid ET production and a programmed cell death at the site of infection that prevents further ingress of the invading pathogen. Several R genes have been identified to confer resistance against insects (Broekgaarden et al., 2011), but whether they are involved in recognizing specific herbivore-derived elicitors is still unclear.
Subsequent to recognition of the attacker, a hormone-regulated cellular signaling network is triggered that orchestrates the production of defensive proteins and metabolites. Besides ET, several other plant hormones are implicated in this regulatory network, with jasmonic acid (JA), salicylic acid (SA), and abscisic acid (ABA) being major players (Fig. 1; Pieterse et al., 2012). Their abundant antagonistic or synergistic interactions provide the plant with an extensive regulatory potential that enables the activation of specific defenses, while minimizing fitness costs (Pieterse et al., 2012; Vos et al., 2013a). ET emerged as an important hormonal traffic controller on the hormone-regulated defense pathways that are activated in response to biotic stress. Here, we provide an update on the modulating role of ET in this process.
During induced plant defenses in general, activation of ET-mediated defenses starts with the recognition of the attacker, after which calcium fluxes and MAPK cascades are initiated that in turn stimulate ET biosynthesis. The amount and timing of ET production is then further regulated in an attacker-specific manner, which modulates the plant's defense response (Hu et al., 2011; Van der Ent and Pieterse, 2012; Groen and Whiteman, 2014; Rehrig et al., 2014). In addition to its role in defense, ET is also known to be an important regulator of developmental processes, and accordingly, ET-dependent defense patterns change during development. For instance, treating Nicotiana attenuata leaves of various ages with saliva from the Manduca sexta caterpillar showed that induced ET emission decreased with leaf and plant age (Diezel et al., 2011).
Recognition of Attack Leading to ET Signaling
A classic example of attacker perception leading to the activation of ET signaling is the recognition of MAMPs such as flagellin and elongation factor (EF)-TU (Boller and Felix, 2009). Upon perception of these MAMPs by their cognate PRRs, MAPK signaling is activated, leading to rapid production of ET, which results in effective downstream PTI responses. In the case of flagellin perception, ET signaling regulates accumulation of the cognate PRR receptor FLAGELLIN SENSITIVE2, and is required for the oxidative burst that contributes to flagellin-triggered plant immunity (Mersmann et al., 2010). Furthermore, plant elicitor peptide ZmPep3 in maize (Zea mays) serves as an endogenous signal to stimulate ET production and increase the expression of genes involved in direct and indirect defense, leading to enhanced resistance against Spodoptera exigua caterpillars (Wu and Baldwin, 2010; Huffaker et al., 2013). A recent example of effector-triggered activation of ET signaling comes from the interaction between the bacterial pathogen Erwinia amylovora and Arabidopsis (Arabidopsis thaliana). The E. amylovora-derived elicitor HrpNEa activates ET-mediated expression of the Arabidopsis transcription factor MYB44, which in turn enhances the expression of ETHYLENE INSENSITIVE2 (EIN2), an essential gene in ET signaling (Liu et al., 2011a).
Molecular Players in Signal Transduction from Recognition to ET Biosynthesis
Recently, several unique components involved in the steps between pathogen/insect recognition and the activation of ET signaling have been reported. For example, during infection of Arabidopsis by the necrotrophic pathogen Botrytis cinerea, MITOGEN-ACTIVATED PROTEIN KINASE3 (MPK3) and MPK6 phosphorylate the ET biosynthesis proteins 1-AMINOCYCLOPROPANE-1-CARBOXYLIC ACID (ACC) SYNTHASE2 (ACS2) and ACS6, resulting in rapid ET production (Han et al., 2010). Additionally, silencing SALICYLIC ACID-INDUCED PROTEIN KINASE (SIPK), encoding the N. attenuata ortholog of Arabidopsis MPK6, abolished insect-induced ET accumulation in N. attenuata (Wu et al., 2007; Mase et al., 2012). Moreover, SIPK- and WOUND INDUCED PROTEIN KINASE-silenced Nicotiana umbratica plants showed disrupted ET accumulation and compromised necrosis upon application of the pathogen-derived Alternaria alternata sp. lycopersici toxin (Mase et al., 2012). Taken together, these results highlight the importance of MAPK signaling cascades in the regulation of ET signaling and its downstream defense responses. Furthermore, the Ca2+-binding protein Calreticulin 3a of Nicotiana benthamiana was shown to be involved in the induced production of ET upon recognition of MAMPs derived from the pathogen Phytophthora infestans. Induced ET signaling was further demonstrated to be required for the production of antimicrobial phytoalexins, which is critical for the resistance of N. benthamiana to P. infestans (Matsukawa et al., 2013).
ET-Mediated Downstream Responses
Downstream in the ET signaling cascade, transcription factors of the ETHYLENE RESPONSE FACTOR (ERF) family play a dominant role in the regulation of defenses. For example, ERF3 was identified in rice (Oryza sativa) as a gene that positively affects expression of trypsin proteinase inhibitors and to mediate resistance toward Chilo suppressalis caterpillars (Lu et al., 2011). Well-characterized ERF transcription factors from Arabidopsis with a role in ET-dependent defense are ERF1 (Lorenzo et al., 2003) and OCTADECANOID-RESPONSIVE ARABIDOPSIS APETALA2/ ETHYLENE RESPONSE FACTOR DOMAIN PROTEIN59 (ORA59; Pré et al., 2008). Overexpressing or silencing these ERF transcription factors in Arabidopsis leads to enhanced or reduced resistance to several fungi, respectively (Memelink, 2009). Overall, ERFs are conserved among different plant species (Groen and Whiteman, 2014) and mediate resistance to especially necrotrophic pathogens (Anderson et al., 2010; Liu et al., 2011b; Zhu et al., 2014).
The ETI response that is triggered in Arabidopsis upon infection by avirulent Pseudomonas syringae pv tomato avrRpm1 was reported to be accompanied by a biphasic accumulation of ET (Mur et al., 2009). This ETI-associated ET production is dependent on ETHYLENE RECEPTOR1 (ETR1) and ET signaling component EIN2 and accelerates the hypersensitive cell death response, confirming previous findings that ET plays a role in lesion expansion during the hypersensitive response (Van Loon et al., 2006).
A plant's response to insect or pathogen attack is based on a complex network of interactions between different hormonal signals. Cross communication between the initiated hormone signaling pathways contributes to the activation of attacker-specific defenses (Robert-Seilaniantz et al., 2011; Pieterse et al., 2012; León et al., 2014; Caarls et al., 2015; Fig. 1). JA and SA are dominant players in the regulation of the immune signaling network, where JA is generally effective against necrotrophic pathogens and insects while SA is primarily effective against biotrophic pathogens (Vos et al., 2013a). Accumulating evidence indicates that ET can interact both positively and negatively with SA, depending on the plant-attacker interaction (Van der Ent and Pieterse, 2012). For example, ET acts positively on the level of SA-mediated resistance against the pathogen Leptosphaeria maculans in Brassica napus (Sašek et al., 2012), while it acts negatively on SA-mediated defense against the pathogen P. syringae in Arabidopsis (Chen et al., 2009). The majority of studies on the modulating role of ET report on its effect on the JA signaling pathway and on the interplay between the JA and SA pathways. Therefore, we focus on these ET-hormone interactions in more detail below.
Interaction between ET and JA Signaling
Although the JA pathway has a dominant role in induced plant defenses (Wasternack, 2015), ET plays an important part in fine tuning these responses. The JA signaling pathway comprises two separate branches: (1) the so-called ERF branch, which in Arabidopsis is regulated by ERF-type transcription factors such as ERF1 and ORA59 (Lorenzo et al., 2003; McGrath et al., 2005; Pré et al., 2008), and (2) the so-called MYC branch, which in Arabidopsis is regulated by MYC-type transcription factors such as MYC2, 3, and 4 (Boter et al., 2004; Dombrecht et al., 2007; Fernández-Calvo et al., 2011). ET synergizes the ERF branch to activate defense-related genes, such as PLANT DEFENSIN1.2 (PDF1.2), leading to effective defense against necrotrophic pathogens. The MYC branch, on the other hand, is synergized by ABA to activate defense-related genes, such as VEGETATIVE STORAGE PROTEIN2, resulting in defense against chewing insects (Fig. 1). The ERF and MYC branches of the JA pathway act as communicating vessels in which the relative balance is dependent on the relative strength of the concomitantly activated ET and ABA pathways (Lorenzo et al., 2004; Verhage et al., 2011; Vos et al., 2013b). This is exemplified by the fact that ABA-deficient mutants are more resistant to necrotrophic pathogens and more susceptible to certain insects, whereas ET-deficient mutants are more susceptible to necrotrophs and more resistant to certain insects (Berrocal-Lobo et al., 2002; Lorenzo et al., 2003, 2004; Nickstadt et al., 2004; van Loon et al., 2006; Bodenhausen and Reymond, 2007; Kazan and Manners, 2012; Dinh et al., 2013). However, it should be noted that there are exceptions; for example, an ET-insensitive mutant of N. attenuata displayed reduced JA-mediated defenses against M. sexta caterpillars (Onkokesung et al., 2010).
Recent studies provide evidence that the balance between the ERF and MYC branch of the JA pathway is modulated through interactions between JA/ABA-activated MYC2/3/4 and ET-stabilized EIN3 and ETHYLENE INSENSITIVE3-LIKE1 (EIL1; Fig. 2). Zhang et al. (2014) showed that MYC2 represses EIN3 by inducing expression of the EIN3-repressor ETHYLENE INSENSITIVE3 BINDING F-BOX PROTEIN1 as well as by physically interacting with EIN3 to inhibit its DNA binding activity. Additionally, Song et al. (2014) showed that this antagonistic effect of MYC2 on EIN3 also involves EIL1 and attenuates resistance to B. cinerea. Furthermore, they demonstrated that the interaction between MYC2 and EIN3 is mutually antagonistic. EIN3 and EIL1 interact with and repress MYC2, MYC3, and MYC4, resulting in attenuated JA-regulated defense against Spodoptera littoralis and S. exigua caterpillars. Important modulators of the activity of the MYC2/3/4 and EIN3/EIL1 transcription factors are JAZ repressor proteins that physically bind to them (Fig. 2B; Chini et al., 2007; Fernández-Calvo et al., 2011; Niu et al., 2011b; Zhu et al., 2011). Another modulator of the activity of the MYC and ERF branches of the JA pathway is the mediator complex subunit MEDIATOR25 (MED25; Çevik et al., 2012). In eukaryotes, the multiprotein mediator complex connects transcription factors to the core transcriptional machinery. MED25 binds to and is required for the transcriptional activation of the MYC and ERF branch transcription factors MYC2, ERF1, and ORA59 (Fig. 2B). Consequently, med25 mutants are impaired in ET- and JA-dependent defenses against pathogens and insects (Çevik et al., 2012).
Effect of ET on SA-JA Cross Talk
Among the best-studied hormone signal interactions is the antagonistic relationship between the SA and JA signaling pathways, often referred to as SA-JA cross talk (Pieterse et al., 2012). A number of important regulators of the interaction between the SA and JA pathways have been identified, including the redox sensitive transcriptional coregulator NONEXPRESSOR OF PATHOGENESIS-RELATED PROTEINS1 (NPR1; Spoel et al., 2003). Interestingly, timing and concentration of ET production modulate the strength of the antagonistic effect of SA on the JA pathway and the dependency on NPR1. Leon-Reyes et al. (2009) demonstrated that simultaneous stimulation of the ET and JA pathways in Arabidopsis strongly suppressed the NPR1 dependency of SA-JA cross talk. Additionally, full activation of ET and JA signaling (e.g. during infection with a necrotrophic pathogen) resulted in insensitivity to future SA-mediated suppression of JA/ET-dependent defenses. Hence, ET helps the plant to prioritize the JA/ET pathway over the SA/NPR1 pathway during multiattacker interactions (Leon-Reyes et al., 2010). Also, in other plant species, a link between ET and NPR1-dependent regulation of defense has been reported. For instance, antisense expression of NPR1 in rice increased JA and ET levels, resulting in increased levels of antiherbivore compounds and reduced performance of rice striped corn borer (Li et al., 2013).
Besides NPR1, other SA-regulated signaling components emerged as important regulators of the interplay between the SA and JA/ET pathways (Fig. 2C). In Arabidopsis, TGA transcription factors are primarily known as regulators of SA-mediated transcription, but they control activity of the ET/JA-dependent ERF branch as well. TGA2, TGA5, and TGA6 are required for B. cinerea-induced expression of PDF1.2 to confer resistance against this necrotrophic fungus (Zander et al., 2010). However, in the presence of SA, the same TGAs have a role in SA-mediated antagonism of ERF branch signaling. The TGAs can directly target the activation sequence-1 (as-1) promoter sequence in the ERF transcription factor gene ORA59, thereby regulating its expression (Zander et al., 2014). Possibly through SA-induced glutaredoxins such as GLUTAREDOXIN480 (GRX480) that interact with TGA transcription factors, expression of ORA59 is suppressed (Zander et al., 2012; Caarls et al., 2015). Together with the observation that SA affects ORA59 protein accumulation (Van der Does et al., 2013), these findings demonstrate that ORA59 plays a central role in the interaction between the SA and JA/ET pathways.
Plants activate defenses not only locally at the site of attack, but also systemically throughout the plant to protect the still-healthy tissue against future attack. Pathogen-induced systemic acquired resistance (SAR), beneficial microbe-induced systemic resistance (ISR), and herbivore/wound-induced systemic resistance are three well-characterized types of systemic defense responses (Wu and Baldwin, 2010; Pieterse et al., 2014b). Although ET seems to have only a minor role in systemic acquired resistance (Fu and Dong, 2013) and herbivore/wound-induced systemic resistance (Vos et al., 2013b), its essential role in the regulation of ISR is widely accepted (Pieterse et al., 2014b). Interestingly, ET is also implicated in volatile signaling, either by serving as a volatile itself or regulating the production of other volatiles to communicate with distant plant tissues or neighboring plants about pathogen or insect attack. As part of an indirect defense mechanism, volatiles also function to repel insects or attract the insect's natural enemies (Scala et al., 2013).
The Role of ET in ISR
Plants produce exudates and lysates at their root surface, resulting in the attraction of a large number of beneficial microorganisms (Berendsen et al., 2012). Besides stimulating plant growth and outcompeting soil-borne pathogens, some beneficial microbes are capable of activating ISR in above- and belowground plant tissues, a phenomenon that is effective against a broad spectrum of pathogens and insects (Pineda et al., 2010; Pieterse et al., 2014b; Zamioudis et al., 2015). In many cases, ET has been demonstrated to play an important role in the modulation of host immune responses by beneficial microbes (Zamioudis et al., 2012). At the root-microbe interface, the MAMP flagellin of the ISR-inducing rhizobacterium Pseudomonas fluorescens WCS417 (recently renamed as Pseudomonas simiae WCS417; Berendsen et al., 2015) initially triggers an ET-dependent immune response in Arabidopsis roots, but this becomes readily suppressed, probably to facilitate accommodation of the microbe on the root surface (Millet et al., 2010). In contrast, activation of ET signaling is essential in the initial stages of colonization of barley (Hordeum vulgare) roots by the mutualistic fungus Piriformospora indica (Khatabi et al., 2012). In the interaction between Populus trichocarpa and the ectomycorrhizal fungus Laccaria bicolor, ET acts together with JA in constraining fungal growth, possibly to maintain an economic balance between costs and benefits in this mutualistic plant-fungus interaction (Plett et al., 2014).
To develop ISR, plants need an intact ET response, as was demonstrated by the impaired ability of ET signaling mutants to develop ISR (Van Wees et al., 2008; Niu et al., 2011a; Salas-Marina et al., 2011; Fracetto et al., 2013; Pieterse et al., 2014b). In tomato (Solanum lycopersicum), the ET dependency of ISR triggered by a mutualistic Methylobacterium spp. strain is related to the suppression of ET accumulation induced by the pathogen Xanthomonas campestris, resulting in a reduction of disease severity (Yim et al., 2014). Hence, both locally in the root and systemically in the leaves, ET modulates immune responses, either to accommodate mutualists or to counteract enemies.
The Role of ET in Volatile Signaling
Besides playing a vital role as endogenous plant hormone, ET is also involved in volatile signaling, especially during plant-insect interactions. ET is not only volatile by itself but also positively regulates the accumulation of volatile organic compounds, often together with JA (Fig. 1; Pierik et al., 2014). For example, exogenous application of the ET precursor ACC on lima bean (Phaseolus lunatus) leaves enhanced the production of three JA-mediated volatiles [i.e. (E)- and (Z)-β-ocimene and (Z)-3-hexenyl acetate], leading to the enhanced attraction of the carnivorous mite Phytoseiulus persimilis (Horiuchi et al., 2001). ET-mediated volatile production can alter insect behavior in an insect species-dependent manner. For example, silencing of ET biosynthesis in rice reduced the volatile release upon infestation by C. suppressalis caterpillars, leading to reduced plant resistance. In contrast, infestation of ET-silenced rice plants by the phloem-feeding brown plant hopper (Nilaparvata lugens) increased the emission of volatiles that repelled this insect (Lu et al., 2014). Besides altering insect behavior directly, ET-mediated volatile production can also play a role in indirect defense through the attraction of carnivorous enemies of the insect (Scala et al., 2013). For example, the S. exigua caterpillar-induced production of JA/ET-mediated volatile terpenes and benzoxazinoids in maize enhanced the attraction of the parasitoid Cotesia merginiventris, which is a natural enemy of S. exigua (Huffaker et al., 2013).
Finally, ET also plays a role in plant-plant communication, in which an attacked plant transfers volatile information to its neighboring plants, which in turn tailor their defenses (Holopainen and Blande, 2012). For example, exposure of intact maize plants to (Z)-3-hezen-1-ol, a volatile emitted by green plants upon mechanical damage, induced the emission of a volatile blend that is typically emitted after caterpillar feeding and attracts natural enemies of the herbivores. The volatile emission increased when ET was added, indicating a synergistic role for ET in plant-plant signaling mediated by (Z)-3-hezen-1-ol (Ruther and Kleier, 2005). Although there is a clear role for ET in plant volatile signaling, knowledge of the function of ET in plant-insect and plant-plant communication is still limited.
ET controls multiple aspects of plant defense by connecting different hormonal signaling pathways and often decisively modulating their relative output. This central role of ET makes it a valid target for attackers in their battle against induced plant defenses. Certain pathogens and insects activate diverse hormone signal integration mechanisms to manipulate the plant's defense system, resulting in the circumvention or suppression of effectual defenses (Walling, 2008; Robert-Seilaniantz et al., 2011; Pieterse et al., 2012; Kazan and Lyons, 2014). ET plays a role in this decoy strategy of several attackers, resulting in increased plant susceptibility. For example, the soil-borne pathogen Fusarium oxysporum produces ET itself, leading to expression of the ET receptor gene ETR1 in Arabidopsis, which in turn leads to suppression of effective SA-dependent defenses, and thus promotion of Fusarium spp. disease (Pantelides et al., 2013). Additionally, saliva of Pieris rapae caterpillars activated the ERF branch of the JA signaling pathway in Arabidopsis, leading to suppression of effective defense responses that are mediated by the MYC branch (Verhage et al., 2011). Moreover, saliva of S. exigua caterpillars increased cellular oxidative stress in an ET-dependent manner in leaves of Medicago truncatula, which led to SA/NPR1-mediated suppression of JA-related defensive proteins (Paudel and Bede, 2015). This suggests that S. exigua engages the plant's ET signaling pathway to activate SA-mediated antagonism of JA signaling for its own benefit. The role of ET in plant defense manipulation by attackers is still poorly understood, and future research in this direction may provide us with novel insight in regulatory components of hormone signal integration.
In past decades, the pathways of ET biosynthesis and signaling have been elucidated in detail. Moreover, ET emerged as an important modulator of other hormonal signaling pathways that play a role in the regulation of plant growth and adaptive responses to biotic and abiotic stresses (Kazan, 2015; Vos et al., 2015). More and more molecular players active at the crossroad of hormone signaling pathways that regulate both growth and defense were identified (Pieterse et al., 2014a; Caarls et al., 2015). Knowledge of master regulators and their gene regulatory networks provides important tools to investigate how the plant's stress signaling network functions during different environmental conditions. In natural and agricultural settings, plants often have to cope with multiple biotic and abiotic stresses at the same time. So how do ET and other hormones steer the plant stress signaling network under these conditions of combinatorial stress? And how can we utilize this knowledge to improve our crops? To obtain a deep understanding of the dynamics of the hormone signaling network, it will be essential to investigate the dynamics of the transcriptome and proteome during single and multistress interactions. Computational analysis of high-density time series of transcriptome changes during ET-inducing stress conditions, such as leaf senescence (Breeze et al., 2011) or infection by the necrotrophic pathogen B. cinerea (Windram et al., 2012), or during cross talk between defense hormones (Van Verk et al., 2013) has been shown to be highly instrumental in the prediction and validation of unique players in the network. Unique technologies such as ChIP-seq (Chang et al., 2013) and protein interactome mapping (Braun et al., 2013) will further integrate ET signaling proteins and transcription factors in the dynamic hormone-mediated signaling network. These systems' approaches will ultimately lead to a better understanding of how the plant growth regulator ET evolved as a traffic controller on the hormonal crossroads to defense.
We thank Roeland Berendsen, Silvia Proietti, and two anonymous reviewers for valuable comments on the article.
↵1 This work was supported by the Dutch Technology Foundation STW, part of the Netherlands Organization of Scientific Research (VENI grant no. 13087 to C.B. and VIDI grant no. 11281 to S.C.M.V.W.), and the European Research Council (ERC advanced grant no. 269072 to C.M.J.P.).
- pattern recognition receptor
- microbe-associated molecular pattern
- pattern-triggered immunity
- ethylene
- mitogen-activated protein kinase
- effector-triggered immunity
- jasmonic acid
- salicylic acid
- abscisic acid
- jasmonic acid-Zim domain
- microbe-induced systemic resistance
- Received July 3, 2015.
- Accepted October 19, 2015.
- Published October 19, 2015.


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Keeping your hormones balanced and happy during the holidays can feel tough during this time of year. The stress, family expectations, and traditionally unhealthy food can throw a wrench in your health plans.
Being constantly on the go and having to travel more than usual to see family can cause the stress-hormone cortisol to spike, leaving you feeling constantly exhausted. On top of that, many of the foods found on holiday dinner tables contain gluten, dairy and refined sugars, which can do a number on your gut (many important hormones, such as serotonin and melatonin, are made in your gut, so if this is not working properly it has a negative effect on all your hormones).
Stress and poor eating can create a snowball effect and end up throwing off your estrogen and progesterone levels – further impacting your gut and digestion for the worse. These changes in your body can manifest in not only fatigue and craving for high sugar food, but cause mood swings, anxiety, bloating, headaches, and can even throw off your menstrual cycle.
Ok, so what can you do about it?
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Eat healthier alternatives.
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Posted on June 23 2019
Beautiful Katy wore the Tanita, one of our simple yet gorgeous silk dresses, to her farm wedding filled with joy.
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I wanted something that felt vintage-y, but not like a costume. I wanted to be able to move easily (and dance!) in the dress. And I needed something that would flatter my boobs. But I apart from that, I had no idea!
The experience of purchasing the dress was really lovely - my first visit was with my bridesmaids, and it just flew by! We were really well looked after. The only difficulty was that I loved everything in the shop, so narrowing it down was really hard. So I came back a second time with my mum to make the final choice, which was also lovely.
I really loved the Tanita dress straight away, but wasn't sure about the train (I was worried about someone stepping on it on the dance floor). I was really pleased that Miina was able to make me one without a train - thank you so much!
Makeup and hair
My makeup and hair was done by Lipstick & Curls.
None really! I wore Swedish Hasbeen shoes, but no other accessories.
Suit, shirt and tie from Reiss. We actually forgot to buy him new shoes for the wedding, so he wore a pair he's had for years instead.
We got married in Upton Cheyney, near Bristol. Wedding ceremony was in the village chapel, and the reception was at Manor Farm.
We didn't have a particular theme. We really wanted to have a relaxed and fun day, so didn't want the pressure of planning something where it felt that everything had to match. So I suppose a loose theme was 'stuff we like'... and so we went with lots of flowers and fairy lights.
The flowers were by Lucy Coco Flowers (Bristol) - such a lovely girl, and really gorgeous flowers! Again, very informal and unfussy.
Food was provided by Manor Farm - bbq for the main meal, and then a ploughmans in the evening. All using meat from the farm and local fruit/veg!
Best Men DJs, and Brass Junkies brass band.
Joanna Nicole Photography - so recommended! We loved every minute of working with her, she made us feel comfortable right and seemed to know exactly what we wanted without us even having to ask.
- Katy -


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|Based on cycle analysis, the Russell 2000 index is anticipated to bottom out (on the daily time frame) around October 7–12, 2009. Can we also calculate a possible price level at which to expect that reversal, should it arrive as expected? The answer is a qualified "maybe."
|FIGURE 1: RUT, DAILY. The Russell 2000 index may bottom on its daily cycle by October 7–12, 2009, possibly between 572 and 565.
|Graphic provided by: MetaStock.
|Graphic provided by: WB Detrend EOD from ProfitTrader for MetaStock.
|Figure 1 is the daily chart of the R2K; the index has had a modest retracement move, one lasting a week and a half — so far. Note how this correction has unfolded in a series of small waves, unlike the prior two corrections, both of which were simple pullback-type moves against a strongly rising weekly price cycle. This latest one is deeper, more convoluted, and is more likely than not to test the next set of daily Fibonacci support levels that span the range from 572 to 565. One reason to believe that lower prices are due is that the internal strength readings of the broad market indexes are lower now that they were at the last two swing lows made in August and September, respectively. If this analysis is correct, we may see a briefer selloff (lasting a couple of days) down to the support level prices (572–565), followed by an actual bottoming/reversal confirmation.
There is a modestly bullish omen on the R2K daily graph, one that also suggests the worst of this daily selloff is behind us; the detrend oscillator (bottom panel of chart) has tentatively printed its second straight hidden divergence against the index price. You don't see much written about this phenomenon in the traditional technical analysis texts, but it is a very useful tool when placed in the proper context. The concept is simple, if price keeps rising, putting in successively higher swing lows even as the detrend oscillator (or even a relative strength index [RSI], stochastic, commodity channel index [CCI], or stochRSI indicator) keeps making successively lower indicator lows, you have a valid hidden divergence. There will frequently be a reversal higher in the wake of such a divergence, with the strength typically determined by the size of the divergence. It's a fascinating pattern, one with above-average reliability, and that's why we need to watch what happens if and when the R2K gets near our anticipated time/price reversal zone. Even though the weekly cycle in the R2K is still heading down, a tradable relief rally on the daily chart is also a likely occurrence in the next week or so after the final (projected) washout into the daily cycle low.
|FIGURE 2: RUT, WEEKLY. While the daily cycle is due to bottom soon, the weekly cycle likely has another five to six weeks to go before a meaningful reversal occurs.
|Graphic provided by: MetaStock.
|And now a look at the R2K weekly graph in Figure 2. The double-smoothed stochastic (bottom panel of chart) does a good job of identifying the weekly price cycle in this index, and, right now, it's heading sharply lower with more downside likely to come. The current momentum of the double stochastics is more bearish than that of the previous drop into the July 10, 2009, cycle low, and that suggests that this cycle (currently 13 weeks old) will likely run the full 18 to 19 weeks typically seen in the R2K's weekly price cycle. If that is the case, don't look for a meaningful weekly low until at least mid- to late November 2009. The index does have some solid chart and Fibonacci support ranging from 550 to 520, so as the cycle plays out it should be easier to attempt to pick a more specific date/price to expect a reversal higher.
|So we have two cycles to watch; the daily cycle, which is anticipated to bottom/reverse within a week or so, and the weekly cycle, which still has another five to six weeks to go before the bulls can finally start to prepare for a possible year-end rally. With the quarterly price cycle pointing higher, such a rally shouldn't be ruled out.
|Writer, market consultant
|Linear Trading Systems LLC
|Jacksonville, FL 32217
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|E-mail address:
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The Psychics Guide to Compatibility Tests
Do you believe that a good relationship may have something to do with the compatibility of both individuals in that certain relationship? This compatibility might affect the situations that may take place between you and your partner, or your future partner, as the case maybe. In this case, this is where compatibility tests come to … [Read more…]


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During the fan preview of the upcoming Game of Thrones Live Concert Experience tour, composer Ramin Djawadi (above, far right) ecstatically revealed a few details about the one of a kind event with Entertainment Weekly.
The idea of bringing the music of Westeros on tour was born during a studio session with none other than the brilliant minds behind the series: George R.R. Martin, David Benioff, and D.B. Weiss.
Djawadi reveals:
We've been working on it for a while. So it just seemed now, after season 6, it was such a fantastic season, so we thought there's a lot of material now to draw from.
Entertainment Weekly (EW) also describes the conceptual video of the event shown to the media which includes Thrones locations such as King's Landing and Winterfell, a 360 degree stage, and the Iron Throne itself.
He tells EW:
We really want to summarize the show the best we can. There's a lot of different locations and events to cover. If you come and watch this concert, you really get a nice summary and a nice look back on the past seasons.
The fan event in Los Angeles also previewed iconic Game of Thrones live music such as the main theme and The Rains of Castamere.
Djawadi revels in the music of the series, the live orchestra, and what he's most looking forward to playing for fans with Entertainment Weekly!
The symphony tour sounds like a truly unique experience and essential for any Game of Thrones fan. Will you be attending when winter comes?


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Matt Helms is a North Carolina native and 20 year veteran in the food service industry. He is the Head butcher and owner of the Chop Shop Butchery.
Over 20 immersive signature events are featured throughout the weekend! Tours, dinners, parties, foraging and more await.
This inspired, friendly and free-spirited food scene tucked in the mountains of Appalachia has been called "Foodtopia," but the locals just call it home. Plan your trip now.
Access to affordable and healthy food and reducing food waste are important initiatives of the signature event's charitable giving goals.


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Article 121 TFEU
From thinktank
Article 121 (ex Article 99 TEC) reads
- 1. Member States shall regard their economic policies as a matter of common concern and shall coordinate them within the Council, in accordance with the provisions of Article 120.
- 2. The Council shall, on a recommendation from the Commission, formulate a draft for the broad guidelines of the economic policies of the Member States and of the Union, and shall report its findings to the European Council.
- The European Council shall, acting on the basis of the report from the Council, discuss a conclusion on the broad guidelines of the economic policies of the Member States and of the Union.
- On the basis of this conclusion, the Council shall adopt a recommendation setting out these broad guidelines. The Council shall inform the European Parliament of its recommendation.
- 3. In order to ensure closer coordination of economic policies and sustained convergence of the economic performances of the Member States, the Council shall, on the basis of reports submitted by the Commission, monitor economic developments in each of the Member States and in the Union as well as the consistency of economic policies with the broad guidelines referred to in paragraph 2, and regularly carry out an overall assessment.
- For the purpose of this multilateral surveillance, Member States shall forward information to the Commission about important measures taken by them in the field of their economic policy and such other information as they deem necessary.
- 4. Where it is established, under the procedure referred to in paragraph 3, that the economic policies of a Member State are not consistent with the broad guidelines referred to in paragraph 2 or that they risk jeopardising the proper functioning of economic and monetary union, the Commission may address a warning to the Member State concerned. The Council, on a recommendation from the Commission, may address the necessary recommendations to the Member State concerned. The Council may, on a proposal from the Commission, decide to make its recommendations public.
- Within the scope of this paragraph, the Council shall act without taking into account the vote of the member of the Council representing the Member State concerned.
- A qualified majority of the other members of the Council shall be defined in accordance with Article 238(3)(a).
- 5. The President of the Council and the Commission shall report to the European Parliament on the results of multilateral surveillance. The President of the Council may be invited to appear before the competent committee of the European Parliament if the Council has made its recommendations public.
- 6. The European Parliament and the Council, acting by means of regulations in accordance with the ordinary legislative procedure, may adopt detailed rules for the multilateral surveillance procedure referred to in paragraphs 3 and 4.


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Supplying Keto/LCHF needs all over Australia. FREE DELIVERY ON ORDERS OVER $119! All other orders $9.95 Shipping (Australia only).
Sale price
Regular price
These cookies have only two words: Indulgent yumminess!A chocolate lover's ultimate snack.
Being grain free and sugar free means that they are okay to have when following a low carb or keto diet - just don't get carried away...
Six tasty cookies to hit your cookie craving!


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14 Aug "Goodnight My Love" by Jesse Belvin
Jesse Belvin was a huge musical inspiration to me.
Jesse Belvin was a huge musical inspiration to me.
I both love and hate books that don't leave a discrete ending for the reader. Have you ever felt the need to write sequels?
I am in the process of finishing the sequel of my book "Up From Where We've Come." I realized early on that it would be impossible to cram my whole life into one volume, so I decided to break it up into multiple books.
There's always another book in the pipeline to write… Tell me about it! Does it have a working title?
I am still early in the process, working on the first draft of the next installment, so I am not interested in divulging my next title, at least not yet.
Some advice other writers have given is that your first book is best sitting in a drawer for a while, because then you feel stronger about chopping up 'your baby'. Do you still have a copy of your first draft? How different is this from the final published version of "Up From Where We've Come"?
I do still have a copy of the first draft. The difference is that the first draft has many more words, because once I get everything written down I have to make sure my writing is as concise as possible. The chopping up and process of elimination is very important to the final book.
Do you have a dedicated writing space? How does it meet your writing needs?
I write when and wherever the spirit hits me, but more often than not at my personal computer. Sometimes I will also write on the couch or in bed. The most important thing, though, is that I am comfortable, no matter where I am writing that day.
What is your writing process? Have you ever thought about changing it? Other authors I have interviewed talk about having an outline – post-it notes in an office, or writing in paper journals. Is there something like that in your writing technique? Or is it all digital for you?
I wrote my first draft of "Up From Where We've Come" on a typewriter, since I started writing the book before I owned my first computer. It was only later that I started typing it into a digital format. Because I am not a trained author and am writing my life story, it is straight from my memory bank instead of using outlines.
How do you know when a book or short story is finished? How do you know to step away and let the story speak for itself?
I am not a trained author so I simply go by rote. I simply follow my inner emotions. Like I said before, I knew I would have a difficult time putting my entire life in one book, so the move from Mississippi to California was a great place to pause for the first volume.
Do you have a preference for e-book or paperback format? This is for both your own reading and your writing.
Since I've never read an e-book, I guess that answer would be quite obvious. I can't speak for an experience I've never had. I have however, read many paper books so, I guess you could call me 'old- school'!
Social media is becoming a big thing. How does managing media outlets come into marketing your brand and your books?
I have a Twitter, and Instagram, and a Facebook, but I am so busy and do not usually have time to look after my social media profiles. I'm afraid I have to leave that up to the experts.
You have answered other sets of interview questions, is there something you wish someone would have asked you? Or conversely, something you wish they hadn't asked?
No not really. Thanks for your time Rosemarie
"Up From Where We've Come" now available at Amazon and My Store
It's worth at this point detailing some of background to Wright's book. Born in 1940 in the Deep South, Wright grew up in an America segregated by race. Although the Union had won the Civil War the century previously, a war fought over the slave system that had made Southern white landowners rich, and the slaves were supposedly given their freedom after emancipation, for many very little had changed by the time Wright was born. Slavery was replaced by another form of servitude, namely sharecropping. Now, instead of slaves being owned by the landowners, the landowners would provide some of their land to the newly 'freed' blacks, as well as poor white farmers, along with some rudimentary housing, equipment and maybe even a mule; a local merchant would provide the farmer with credit to purchase food and supplies. In return, at harvest the sharecropper would take a share of the crop produced; the sharecropper would take the rest, minus anything else was owed.
One of the stories that stand out in the book is the first time that Wright took a ride in a car into Clarksdale with the rest of his family. His amazement at being in a car for the first time is fascinating to read for a modern audience who've been around cars all their lives. Yet their ownership of the car did not last long: Wright's father had bought the car with the financial help of Mr Miles, but further financial woes and the devious assistant to Mr Brookings, a man named Leo, conned Wright into selling the car, a blow to Wright's father who tried for years to better his family's desperate situation.
Check out this clip of my live performance at The Saban Theater New Years Eve 2015
Music is my passion. Then, Now, Always
Click Here to read the full review


text: Praise: Millions of church young people youth nationwide hoped to raise $4 million on February 3, 2002, as part of the Souper Bowl of Caring. Youths stood at the doors of their home churches with soup pots, asking parishioners to donate $1 to support local charities. The event coincided with the NFL's Super Bowl. "The Souper Bowl of Caring demonstrates the power of working together," said Brad Smith, Souper Bowl's founder and executive director. "Our nation-wide effort is a subtle reminder-a mustard seed-that ordinary people, with God's help, can do extraordinary good if we work together." In 2001 the drive raised $3.6 million from more than 12,500 congregations from all 50 states, Canada, and Puerto Rico. An estimated 3.5 million participated in the grassroots drive last year.


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Start the weekend early with one of a quartet of special deals and celebrations today. Aurora will be wrangling a herd of food trucks, TAG Burger Bar and Axios are both celebrating birthdays, and wining and dining will even help save some animals. Here's a taste of what's on the culinary calendar...
See also: Beer Calendar: New Brews, a GABF Winner and Pumpkins
TAG Burger Bar will be celebrating its second anniversary with a burger and beer special all day long: $7 gets you a classic burger and a pint of Kannah Creek Blonde.
The Aurora Food Truck Rodeo will be lassoing diners from 11 a.m. to 2 p.m. today at the Aurora Municipal Center, with featured food trucks including Freddy's Latin Cuisine, Manna From Heaven, Mythos Greek and Sweet Cow Ice Cream. Admission is free.
The Pouring for Paws wine-tasting and silent auction event will feature light appetizers, cheeses, live music from Joel Tromburg, and lots of wine. All proceeds will go to save dogs and cats at Outpaws Animal Rescue. The wine will be pouring from 5 to 8 p.m. at the Lumber Baron Inn. Tickets are $60; for tickets and more information on the Outpaws rescue click here.
Axios Estiatorio will be celebrating its third anniversary with live music from San Francisco band Fotia starting at 8 p.m.; admission is $10 at the door. For more information, call Axios at 720-328-2225.
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The more we know about the world around us, the more independent we can be
Do you know how most of the things you use on a daily basis work? Do you know how to fix them when they break?
More often than not, my guess is the answer is: no. And most of us won't even care, because we are abled the luxury to simply replace our defect items. But this comes at a price: dependence.
When me and my partner couldn't take a warm shower during a week in winter because our boiler broke (and the landlord decided to replace it with a new model), it hit us how dependent we were. It was out of our control, given our circumstances. It also hit us how much of a simple warm shower is a luxury we take for granted.
Gone seem to be the days of our ancestors, when if something broke you or someone in your household fixed it themselves. Unless you had to wait for parts, I guess.
We tend to look back on it and see it only as a necessity when they couldn't afford to buy as much as we can now. But maybe that's because consumerism is so pervasive these days, that we can't even see things from a different angle. Consumerism is a very wasteful practice, but it's easy for us to forget about it in our day to day lives because it is convenient and for some of us how we've always done things.
The commodity of consumerism has allowed us to forget and dismiss knowledge and the creativity of how to be independent.
Today I find myself dependent on manufacturers, so called experts and professionals. Not even dependent on people or community, but on a wasteful and impersonal system run by numbers.
Knowledge is a powerful tool, which should be shared.
When knowledge is used wisely, it can be put to the service of all of us and heighten our collective well being. Although, our story so far has been shaped by an unwise application, hoarding information, keeping it secret and letting only a few benefit at the expense of others. Which is clearly visible in our current capitalist system.
But I believe that is something we, as humanity, are already leaving behind us. I believe we are embracing an era of shared knowledge. We have a huge network of open source software and whatnot these days to back this up.
When we learn to do something ourselves, we take back a little of our freedom. We empower ourselves. When we learn about the world around us, we gain the power of knowledge and the ability to act according to our believes, we are no longer ignorant. We are empowered, for we know we have an impact on our surroundings.
Do you know what the story behind your things is?
Do you ever wonder how your things are made? Where they came from? Who made them? Under what conditions? How they made their way to where you bought them from? What happens to them once you throw them away? Where does it go? What happens to it? And how does that impact your world?
I'll be honest, I don't know the answer to most of these questions myself. But I'm learning. I'm learning and I'm starting to make decisions based on that information. Ignorance no longer feels like an option for me.
Back to our cold showers, on that winter week, we turned to the internet and got crafty: we put together a DIY camping shower, where we poured some cold and boiled some water on a tea kettle. Sure it wasn't perfect, but we took back a little piece of ourselves.


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{4.5/5} "When it was over, and he was still trying to catch his breath, the chairman knew that his perceptions had changed again. He'd acquired an appreciation for the power and beauty of the universe that could never be acquired through a mere telephoto lens or a slice of poetry."
Thunderbird by Jack McDevitt, published in 2015
A teleportation device was found on a Sioux reservation, and for now they're controlling access to it. It goes to 3 different places — a world called Eden, a maze of twisty little passages, and a space station outside our galaxy. They're letting scientists slowly explore Eden, when they stumble on a sentient ape-like creature. The teleportation device on Eden leads to a city where short technologically advanced aliens live — possibly the people who built the device in the first place.
This is the sequel to Ancient Shores. You don't have to have read the first one to read this one — but it's worth reading too.
It's about exploration, first contact, and how different people react to unusual events. As usual, McDevitt's novel is highly enjoyable.
I previously reviewed McDevitt's novel Time Travelers Never Die.


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Man dies after motorcycle collides with wild boar in France
PUBLISHED: 14:28 22 March 2019
A 44-year-old man from Nailsea died of multiple injuries after his motorbike hit a wild boar on a motorway in France.
Garry Scutcher, a married accountant from Rowan Close, was found lying on his back on highway A28 at Abbeville-Rouen in France just before 3am on April 27 last year.
Mr Scutcher was discovered by a Jeremy La Forge, a lorry driver, who called the emergency services.
He had suffered multiple injuries to his head, neck, chest, abdomen and pelvis and was pronounced dead at the scene.
In a police interview, which was read out at the inquest of Mr Scutcher at Avon Coroner's Court, in Flax Bourton, on Thursday, Mr La Forge said: "I noticed something that looked like debris in the road.
"I turned my headlights on full beam and I could see a person lying in the middle of the lane.
"I had sufficient time to break and I turned my headlights on and approached the person lying in the road.
"I realised he was dead.
"There was blood in the area of the skull and his helmet must have been ejected as it was no longer in place.
"I also noticed a wild boar and motorcycle in the emergency stopping lane.
"Presumably, he was driving along on his motorcycle and struck a wild boar."
Mr La Forge said he had been droving along the motorway frequently, for a number of years, and had never seen an animal cross the road like that before.
Sergeant Fabrice Desperelle, from Abbeville police station, was called out to the incident.
In a statement read out in court, he said: "When I arrived the emergency services were already there.
"The remains of a wild boar were in the left-hand lane and meters away was a damaged motorcycle.
"Further along from the motorcycle was the body of a person lying on his back and paramedics were performing first aid."
A post-mortem examination by Dr Russell Delaney confirmed Mr Scutcher's injuries were 'unsurvivable'.
He said the nature of his head injury would have rendered him immediately unconscious.
Assistant coroner Terence Moore concluded Mr Scutcher died as a result of a road traffic collision.
If you value what this story gives you, please consider supporting the North Somerset Times. Click the link in the orange box above for details.


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Yubico YubiHSM 2
Ovviamente, è naturale mettere la sicurezza più stretta sui server che memorizzano i segreti custoditi per tutte le autenticazioni degli utenti. Se questi server sono compromessi significa che la sicurezza di tutte le chiavi di crittografia e password residenti su tale server è compromessa – in altre parole, un disastro.
Yubico ha originariamente sviluppato YubiHSM, (un "Hardware Security Module"), per elaborare la codifica, la decodifica e la conservazione dei segreti sui propri server. Oggi, lo YubiHSM2 è l'offerta di Yubico per una facile, conveniente e sicura protezione dei segreti di autenticazione legati alla Yubico OTP memorizzati sul server di autenticazione o di chiavi. Il dispositivo protegge i dati a riposo contro gli attacchi di intrusione condotti a distanza e le minacce interne.
La versione attuale di YubiHSM2 comprende un elemento sicuro e un cambio dal formato originale più grande, verso un design più piccolo (nano) con imbracatura in plastica pressofusa.
Microsoft Active Directory Certificate Services
YubiHSM 2 fornisce una chiave hardware per proteggere le chiavi digitali utilizzate in un'implementazione PKI basata su Microsoft. La distribuzione di YubiHSM 2 ai servizi Microsoft Active Directory Certificate non solo protegge le chiavi root della CA ma protegge anche tutti i servizi di firma e verifica mediante la chiave radice.
Abilita operazioni crittografiche Hardware-Based
YubiHSM 2 può essere utilizzato come toolkit crittografico completo per un'ampia gamma di applicazioni commerciali e open source. Il caso d'uso più comune è la generazione e la verifica della firma digitale basata su hardware.
Migliora la protezione delle chiavi crittografiche
YubiHSM 2 è una soluzione affidabile per la generazione, l'archiviazione e la gestione sicura di chiavi digitali, comprese le funzionalità essenziali per generare, scrivere, firmare, decodificare, hash e wrap keys.
L' hardware security module più piccolo al mondo
Semplice installazione in una porta USB-A
Feature Overview
- Secure key storage and operations
- Extensive cryptographic capabilities: RSA, ECC, ECDSA (ed25519), SHA-2, AES
- Secure session between HSM and application
- Role-based access controls for key management and key usage
- 16 concurrent connections
- Optionally network sharable
- Remote management
- Unique "Nano" form factor, low-power usage
- M of N wrap key Backup and Restore
- Interfaces via YubiHSM KSP, PKCS#11, and native libraries
- Tamper evident Audit Logging
Feature Details
Secure key storage and operations
Create, import, and store keys, then perform all crypto operations in the HSM hardware to prevent theft of keys while at rest or in use. This protects against both logical attacks against the server, such as zero-day exploits or malware, and physical theft of a server or its hard drive.
Extensive cryptographic capabilities
YubiHSM 2 supports hashing, key wrapping, asymmetric signing and decryption operations including advanced signing using ed25519. Attestation is also supported for asymmetric key pairs generated on-device.
Secure session between HSM and application
The integrity and privacy of commands and data in transit between the HSM and applications are protected using a mutually authenticated, integrity and confidentiality protected tunnel.
Role-based access controls for key management and key usage
All cryptographic keys and other objects in the HSM belong to one or more security domains. Access rights are assigned for each authentication key at creation time which allow a specific set of cryptographic or management operations to be performed per security domain. Admins assign rights to authentication keys based on its use case, such as a event monitoring app that needs the ability to read all audit logs in the HSM, or a Registration Authority that needs to issue (sign) end user digital certificates, or a domain security admin who needs to create and delete crypto keys.
16 concurrent connections
Multiple applications can establish sessions with a YubiHSM to perform cryptographic operations. Sessions can be automatically terminated after inactivity or be long-lived to improve performance by eliminating session creation time.
Network Sharable
To increase the flexibility of deployments, the YubiHSM 2 can be made available for use over the network by applications on other servers. This can be especially advantageous on a physical server that is hosting multiple virtual machines.
Remote Management
Easily manage multiple deployed YubiHSMs remotely for the entire enterprise – eliminate on-call staff complexity and travel expense.
Unique "Nano" form factor, low-power usage
The Yubico "Nano" form factor allows the HSM to be inserted completely inside a USB-A port so it's completely concealed – no external parts that protrude out of the server back or front chassis. It uses minimal power, max of 30mA, for cost-savings on your power budget.
M of N wrap key Backup and Restore
Backing up and deploying cryptographic keys on multiple HSMs is a critical component of an enterprise security architecture, but it's a risk to allow a single individual to have that ability. The YubiHSM supports setting M of N rules on the wrap key used to export keys for backup or transport, so that multiple administrators are required to import and decrypt a key to make it usable on additional HSMs. For example in an enterprise, the Active Directory root CA private key might be key wrapped for 7 administrators (M=7) and at least 4 of them (N=4) are required to import and unwrap (decrypt) the key in the new HSM.
Interfaces via YubiHSM KSP, PKCS#11, and native libraries
Crypto enabled applications can leverage the YubiHSM via Yubico's Key Storage Provider (KSP) for Microsoft's CNG or industry-standard PKCS#11. Native libraries are also available on Windows, Linux and macOS to enable more direct interaction with the device's capabilities.
Tamper evident Audit Logging
The YubiHSM internally stores a log of all management and crypto operation events that occur in the device and that log can be exported for monitoring and reporting. Each event (row) in the log is hash chained with the previous row and signed so that it's possible to determine if any events are modified or deleted.
Advanced, Affordable Hardware Security
Whether you are getting started securing servers, or replacing pre-existing Hardware Security Module (HSM) solutions, YubiHSM 2 offers robust capabilities and benefits at a price that is now within easy reach for enterprises.
Trusted Brand
The YubiHSM 2 evolved from the award-winning YubiKey, trusted by 9 of the top 10 internet companies, and 2 of the top 3 financial, retail, healthcare and research institutions globally. Manufactured in USA and Sweden with high security and quality.
Specifiche tecniche
Supporto Sistemi Operativi
|Sistema Operativo
|CentOS 7
Debian 8
Debian 9
Fedora 25
Ubuntu 1404
Ubuntu 1604
|Windows 10
Windows Server 2012
Windows Server 2016
|10.12 Sierra
10.13 High Sierra
Cryptographic interfaces (APIs)
Microsoft CNG (KSP)
PKCS#11 (Windows, Linux, macOS)
Native YubiHSM Core Libraries (C, python)
Cryptographic capabilities
Hashing (used with HMAC and asymmetric signatures)
2048, 3072, and 4096 bit keys
Signing using PKCS#1v1.5 and PSS
Decryption using PKCS#1v1.5 and OAEP
Elliptic Curve Cryptography (ECC)
Curves: secp224r1, secp256r1, secp256k1, secp384r1, secp521r, bp256r1, bp384r1, bp512r1, curve25519
Signing: ECDSA (all except curve25519), EdDSA (curve25519 only)
Decryption: ECDH (all except curve25519)
Key wrap
Import and export using NIST AES-CCM Wrap at 128, 196, and 256 bits
Random numbers
On-chip True Random Number Generator (TRNG) used to seed NIST SP 800-90 AES 256 CTR_DRBG
Asymmetric key pairs generated on-device may be attested using a factory certified attestation key and certificate, or using your own key and certificate imported into the HSM
Performance varies depending on usage. The accompanying Software Development Kit includes performance tools that can be used for additional measurements. Example metrics from an otherwise unoccupied YubiHSM 2:
- RSA-2048-PKCS1-SHA256: ~139ms avg
- RSA-3072-PKCS1-SHA384: ~504ms avg
- RSA-4096-PKCS1-SHA512: ~852ms avg
- ECDSA-P256-SHA256: ~73ms avg
- ECDSA-P384-SHA384: ~120ms avg
- ECDSA-P521-SHA512: ~210ms avg
- EdDSA-25519-32Bytes: ~105ms avg
- EdDSA-25519-64Bytes: ~121ms avg
- EdDSA-25519-128Bytes: ~137ms avg
- EdDSA-25519-256Bytes: ~168ms avg
- EdDSA-25519-512Bytes: ~229ms avg
- EdDSA-25519-1024Bytes: ~353ms avg
- AES-(128|192|256)-CCM-Wrap: ~10ms avg
- HMAC-SHA-(1|256): ~4ms avg
- HMAC-SHA-(384|512): ~243ms avg
Storage capacity
- All data stored as objects. 256 object slots, 128KB (base 10) max total
- Stores up to 127 rsa2048, 93 rsa3072, 68 rsa4096 or 255 of any elliptic curve type, assuming only one authentication key is present
- Object types: Authentication keys (used to establish sessions); asymmetric private keys; opaque binary data objects, e.g. x509 certs; wrap keys; HMAC keys
- Mutual authentication and secure channel between applications and HSM
- M of N unwrap key restore via YubiHSM Setup Tool
Software Development Kit
A Software Development Kit for YubiHSM 2 is available for download on Yubico.com and includes:
- YubiHSM Core Library (libyubihsm) for C, Python
- YubiHSM Shell (Configuration CLI)
- PKCS#11 Module
- YubiKey Key Storage Provider (KSP) for use with Microsoft
- YubiHSM Connector
- YubiHSM Setup Tool
- Documentation and code examples
Physical characteristics
- Form factor: 'nano' designed for confined spaces such as internal USB ports in servers
- Dimensions: 12mm x 13mm x 3.1mm
- Weight: 1 gram
- Current requirements 20mA avg, 30mA max
- USB-A plug connector
Safety and environmental compliance
Host interface
- Universal Serial Bus (USB) 1.x Full Speed (12Mbit/s) Peripheral with bulk interface.


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the effect of revision of a credit rating, when the rating assigned earlier remains unchanged.
the effect of revision of a credit rating, when the rating assigned earlier remains unchanged.
a service furnished by a credit rating agency which is formalized in a resolution of the rating committee, which result is evaluation of creditworthiness of an entity in general by the national rating scale or entity's capacity to pay principal and/or interest on its obligations.
an independent opinion of a credit rating agency on a bank's capacity to timely and in full meet its commitments on deposit obligations.
a result of revision of a credit rating, by which the rating assigned previously, is either upgraded or downgraded.
a system of relations between investors – the company's owners, its managers and other interested parties, which is aimed at balance in influences and interests of participants in corporate relations and at securing company's efficient performance.
an indicator which exhibits risks of transferability and convertibility of foreign currency in the given country. The country ceiling is the highest extreme, which limits individual credit ratings of non-sovereign issuers (companies, banks etc.) in national and foreign currencies. The country ceiling is used by international rating agencies when assigning credit ratings by international scale. By the methodologies of the leading global credit rating agencies, an individual rating of a non-sovereign issuer may be higher than that assigned to the country. However, neither the individual counterparty rating nor the rating of its debt instrument or the country's sovereign rating cannot pierce their country ceiling.
a relative evaluation of general entity's creditworthiness and/or its debt obligation by the national rating scale. The credit rating is accompanied by the rating outlook.
Watchlist is a list of credit ratings being monitored closely by the rating agency for specific rating factors subject to additional analysis that may cause changes of ratings within the period of 6 (six) months.
an aggregate securities of a certain type (of a common face value) issued by a single economic entity; or a loan by which an economic entity carries obligations to repay principal and/or interest in due term, or to provide other proprietary rights, except the rights for managing its own activities.
Payment of interest as well as principal amount on debt obligations of the borrower has been terminated without reaching an agreement with the creditors concerning debt restructure before its maturity.
indicates high probability for the rating's change in the course of the year.
a company, financial institution or other economic entity.
the factors, which maintain or constrain the rating.
the information, which non-disclosure or distortion may have impact on the rating's level.
a written document composed by the rating agency and sent to its customer. It contains the list of facts and of documents necessary for the rating procedure, and the terms within which these facts and documents should be submitted to the agency.
An independent opinion of the rating agency on the insurance company capability of fulfillment obligations on insurance indemnities or cash surrender value refund.
the scale by which international rating agencies assign ratings. The ratings by this scale may be assigned to all obligors and issuers of debt securities irrespectively of their domicile. This enables investors to evaluate credit risks in global prospective and to compare creditworthiness of companies domiciled in different countries. At that, the international credit rating may generally not be higher than the sovereign credit rating of the issuer's domicile. In countries with low sovereign rating (including Ukraine) the possibilities for employment of international rating scales for domestic financial market are limited considerably. A low Ukraine's sovereign rating keeps all Ukrainian obligors within the speculative grade of the international scale. Generally, any of Ukrainian obligors is assigned a low international rating, which makes comparison within a country impossible.
a range of ratings assigned to obligors (or to their debentures), which exhibit higher financial strength and are more reliable in comparison with other obligors (debentures). The investment grade encompasses the long-term credit ratings of uaAAA, uaAA, uaA, uaBBB¸ and the short-term credit ratings of uaK1, uaK2, uaK3.
an independent opinion of a credit rating agency with regard to creditworthiness of certain borrowers towards the debt obligation.
an aggregate of certain type of securities (of a common face value) issued by a single issuer, and which have a uniform type of issue and international identification number, and guarantee their holders equal rights irrespectively of time of purchase or offering.
a legal entity that issued securities and carries obligations to holders of the securities.
an independent opinion of a credit rating agency with regard to creditworthiness of a certain obligor (issuer).
a rating scale for assessment of creditworthiness of obligors and issuers of debt securities within a single country excluding effect of sovereign risks. The possibility for assigning the full range of ratings by the National scale provides wider differentiation of issuers and issues than by the international scale, where ratings generally concentrated at the level lower than the sovereign rating. The national credit ratings are indicated by a special suffix, which denote its country, for instance 'ua' for Ukraine's national ratings.
Both national and international rating agencies may assign national scale ratings. At that the national scale ratings employ the National Rating Scale approved by the resolution of the Government of Ukraine #665 dated Apr. 26, 2007 (http://zakon.rada.gov.ua/cgi-bin/laws/main.cgi?nreg=665-2007-%EF), with each of the international rating agencies operating in Ukraine, employing their own national rating scales. Pursuant to the resolutions of the State Commission for Securities and Stock Market #
On Dec. 27, 2007 the charts of correspondence between national Ukrainian scales of international rating operating in Ukraine and the National Rating Scale approved by the Government of Ukraine.
indicates that there is a possibility to downgrade the rating in the course of the year, on condition that negative tendencies are retained and current risks are realized.
a rating action performed in case the agency discovers factors, which may prompt change of the rating in upcoming 6 (six) months and which require special considerations.
outlook indicates that there is a possibility to upgrade the rating in the course of the year, on the condition that positive tendencies are maintained and that current risks are reduced.
a credit rating assigned with consideration of occurrence/non-occurrence of events which may significantly affect the entity rated. A provisional credit rating may be affirmed or changed following the obtaining supplementary information upon occurrence/non-occurrence of the events.
a document, which contains considerable factors, summary and appendices that the rating report contains and that are free of confidential information on the entity rated.
a conditional sign of the position of the entity relatively to other similar entities.
an action with regard to the entity rated and/or its certain debt obligation as follows:
a collegiate permanent body of a rating agency, which includes agency's experts authorized to make decisions on rating actions.
an independent opinion of a credit rating agency with regard to the company's existing framework of corporate governance, its compliance with the interests of financially interested parties, primarily of its owners. The rating of corporate governance allows for differentiation companies by their quality of corporate governance and with consideration of national peculiarities.
a comment on the prevailing trends, which affect a credit rating in terms of its possible but not mandatory change.
a document, which contains considerable factors, summary, results of the rating research and appendices.
assignment and revision of a credit rating by the National rating Scale basing on contractual relations between the rating agency and a customer.
is performed simultaneously with the rating's change or affirming, which is executed by a resolution of the Rating Committee.
a document which contains data on a credit rating, its outlook, rating action, and date of the resolution.
a service of a rating agency which is formalized by a resolution of the rating committee, which effect is confirming or changing the rating assigned earlier.
is developed by Credit-Rating and intended for determining relative strength of deposits in a certain bank as compared to deposits in other Ukrainian banks.
is developed by Credit-Rating and allows for differentiating companies by quality of their corporate governance.
documents of a due form, which represent pecuniary or other proprietary rights, sets relations of an entity they were offered by, and their holders; they also stipulate fulfillment of obligations on their offering and options for transfering rights to other persons.
a credit rating assigned to obligations of a sovereign country which mirrors capacity and willingness of a sovereign issuer (government) to meet its commitments on the debt obligations in timely and full manner, i.e. to make timely payments on bond issues. This type of rating is also called evaluation of default risks of this country.
a range of ratings which exhibit lower, as compared with the investment grade, financial capacity of an obligor to meet its obligations. The speculative grade includes the long-term ratings of uaBB, uaB, uaCCC, uaCC, uaC and the short term ratings of uaK3, uaK4, uaK5, uaKD.
indicates that there are no anticipated reasons to change the rating in the course of the year
condensed conclusions on the effects of the rating research.
an action of a credit rating agency formalized in a resolution of the rating committee upon which adoption the rating becomes temporarily invalid. The suspended rating may be further revised or withdrawn.
an action of a rating agency which is formalized by the resolution of the rating committee upon which adoption a rating becomes invalid.


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At a ceremony held in New York City on September 19, 2014, leading geneticist and humanitarian Prof. Mary-Claire King, the American Cancer Society Professor of Genome Sciences and Medicine at the University of Washington in Seattle and 2008 Honorary Doctor of Philosophy of Tel Aviv University, was awarded the 2014 Lasker-Koshland Special Achievement Award in Medical Science.
In addition to her honorary degree from TAU, Prof. King was the 2012-2013 Sackler Lecturer at the TAU School of Medicine, and for the past 18 years she has been the research partner of Prof. Karen Avraham, Professor of Human Molecular Genetics and Vice Dean at the Sackler Faculty of Medicine.
The prize, awarded annually since 1946 by the Albert and Mary Lasker Foundation, is considered by many the "American Nobel."
"Prof. Mary-Claire King has employed her intellect, dedication, and ethical sensibilities to generate knowledge that has catalyzed profound changes in health care, and she has applied her expertise to promote justice where nefarious governments have terrorized their citizens," the foundation stated.
Prof. King is one of the world's leading scientists in the field of human molecular genetics. She was the first to demonstrate that a genetic predisposition for breast cancer exists as the result of inherited mutations in the gene she named BRCA1. More recently, she devised a plan to screen for all genes that predispose to breast and ovarian cancers.
Prof. King also pioneered the use of genetic identification to identify victims of human rights violations. Beginning in the 1980s, Prof. King helped find children in Argentina taken from their families during the "Dirty War" of the late 1970s and early 1980s, using a novel approach based on mitochondrial DNA sequencing. She used the same method to identify victims of the bombing of the Jewish community center in Buenos Aires in 1994, and victims of other human rights abuses, natural disasters and man-made tragedies.
For more, visit the Lasker Foundation Web site at http://www.
American Friends of Tel Aviv Universit supports Israel's most influential, most comprehensive, and most sought-after center of higher learning, Tel Aviv University (TAU). A leader in the pan-disciplinary approach to education, TAU is internationally recognized for the scope and groundbreaking nature of its research and scholarship -- attracting world-class faculty and consistently producing cutting-edge work with profound implications for the future.
TAU is Israel's only institution of higher learning ranked among the world's top 200 universities by the authoritative Times Higher Education World University Rankings. It is one of a handful of elite international universities rated as the best producers of successful startups, and TAU alumni rank 9th in the world for the amount of American venture capital they attract.


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The hiring of a law firm is an important decision that should not be based solely on advertisements. Before you decide, ask us to send you free written information about our qualifications and experience. Personal Injury Lawyers serving Daytona, Melbourne, Orlando, Florida and Nationwide.This website is not intended for viewing or usage by European Union citizens.
Posted: August 10, 2013
The U.S. government has earmarked $ 2,000,000 to go to the cities in the nation with the highest pedestrian fatality rates. This money is to be used for pedestrian safety enforcement, education and awareness programs. Already, the National Highway & Traffic Safety Administration has created a website devoted to pedestrian safety and awareness. The NHTSA website can be found at Everyone Is A Pedestrian. The site has tips for parents and caregivers, children and aging adults. There are also guides for teachers, educators, and consumer safety advocates.
According to the NHTSA, over 4400 pedestrians were killed in traffic accidents in 2011. Locally in Central Florida, we all know that Metro Orlando won the dubious distinction of being on the top most dangerous cities in the U.S. for pedestrians. Hopefully, local legislators will be taking advantage of this federal program to bring pedestrian safety awareness to Central Florida as a reality.
If you or a loved one has been injured in an automobile accident, contact us today for a free consultation about your legal rights.


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CNBLUE has released a highlight medley for their upcoming mini album, '7°CN'. This will allow fans a sneak peek into the type of tracks the boys have composed and written. Check it out below!
There is a total of six tracks including the title track, 'Between Us', 'It's You', 'Calling You', 'When I Was Young', 'Manito' and 'Royal Rumble'. Even though it is very short, each track so far already sounds amazing. Each song has their unique beat that is addictive to listen to. Especially the title track shows off the pleasant vocals that runs soothingly to the ears. It definitely has fans excited for its full release! From the medley, which track caught your attention the most? Let us know in the comments below!
CNBLUE has also released the band performance trailer for their title track, 'Between Us'. It shows off the amazing skills that CNBLUE has. Especially, when Minhyuk is concentrating hard on playing the drums and bass, Jungshin's smile while playing the guitar, Jonghyun riding the beats with his guitar and Yonghwa passionately singing the chorus, it just proves to the fans how much this means to CNBLUE!
The boys will be back on the 20th of March. Meanwhile check out the highlight medley and the band performance trailer above! Did you like their tracks so far? Are you excited for their comeback? Let us know in the comments below!
Officially Kmusic will continue to keep you updated on Korean music, so stay tuned!
Source: CNBLUE Youtube Channel
Written and edited by: Tracey
- Excited
- Fascinated
- Amused
- Bored
- Sad
- Angry


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The MicroChem Kit 2nd Edition contains sufficient materials to perform every experiment five times with all that's needed except a few commonly available items. It is intended as full year of labs designed to make teaching and preparation very convenient for the instructor. Use with adult supervision. Ages 14 & up.
Lab manual with full instructions is included in the lab kit.
Lab Experiments:
- Scientific Method
- Paper Chromatography
- Collecting Data
- Atomic Orbital Models
- Modeling Carbonate Reactions
- Electrical Conductivity
- Hybridization of Orbitals
- Decomposition
- Double Replacement Reaction
- Analysis of Hydrates
- Mole Ratios
- Boyle's Law
- Charles's Law
- Melting Points
- Freezing Point Depression
- Enthalpy of Ice
- Reversible Reactions
- Reaction Rates, Concentration
- Reaction Rates, Temperature
- Solubility Product Constant
- pH and pH Indicators
- Titration
- Molar Mass by Titration
- Buffers
- Oxidation-Reduction
- Galvanic Cells
- Organic Chemistry Models
- Hydrocarbon Models
- Polymer Models
- Cross-linking of a Polymer
- Nuclear Decay Simulation
29 Chemicals, 27 different types of equipment, 200 page manual, 60 page teachers answer guide – all in a convenient storage box. Developed with safety in mind, the kit contains no concentrated acids or bases. The equipment is mostly plastic to reduce breakage.
Note: Lab kits are not available to ship outside the United States (except Canada and Mexico) as they can only ship via ground service. You can most likely find a Chemistry kit supplier in your country that has similar components as this is an off the shelf lab kit. You can order the other Oak Meadow Chemistry items a la carte.


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Punjab And Haryana Court SSSC Clerk Result 2019 : The high court of Punjab & Haryana is conducted subordinate staff clerk examination 2019 on 04th August. More than 2 lakhs hopefuls were join the written test on various exam center in both state. Department organize competition test for Advt. No. 25C/SSSC/HR/2019. HC of P&H at Chandigarh is proceeding this recrement to hire valuable contenders on Clerk Position. Eligible people with graduation degree and Minimum 18 Yrs. to maximum 42 Yrs. Appear in the examination. All of them are requiring any basic information like expected Cut off marks of Punjab and Haryana high court SSC Clerk Exam 4/8/2019. To login for such information you may have to secure your admit card / roll number and download High Court Punjab & Haryana Clerk 2019 Results online.
Punjab And Haryana Court SSSC Clerk Result 2019 – Chandigarh HC Clerk Cut off Marks / Merit List
Major test of elimination and selection students is finally done by HC of Chandigarh, Punabj & Haryana HC. Under its official portal www.sssc.gov.in is shortly upload selection of students file as PDF. In August / Sept. month tentatively dates are fixed dynamically by HR Team. Eager students for download Punjab & Haryana High Court Clerk Written Marks Details 2019 examination. A new schedule is going to soon that will fix such issues of when will declare HC Punjab Haryana Clerk Exam Results 2019. The direct link if HC PH Clerk august Exam 2019 is mention on this page whenever department publish it.
Punjab and Haryana HC Clerk Results Date 2019
An Overview
|Subordinate Staff Clerk Recruitment 2019
|SSSC Clerk
|No. of Posts
|352 Vacancies
|Exam Day
|07/01/2019 to 04/02/2019
|Article Type
|Official Link
|Selection Method
|Written + Typing Test
|Result Date
|Announce Soon
About Exam & Selection :
Written Test : SSSC Clerk Written Exam 2019 was contains question paper with 50 Questions of General Knowledge and 50 Questions of English Composition. Minimum 33% marks is required as minimum qualifying marks and 40% marks in the aggregate in both subjects. Results will prepare after calculating ¼ marks as deduct for each wrong attempted answer by all students. After that score marks is announce to you all guys.
Typing Test : the Central Recruitment Committee will called shortlisted students for computer proficiency test if they are qualified in written examination. CPT is comprise two parts as Part-I word processing test, English typing on computer with Speed of 30 W.P.M. and In Part-II you will examinee for Spread sheet test knowledge.
Download Haryana & Punjab HC Clerk Cut-off Marks / Merit List
Haryana And Punjab High Court under Society for Centralized Recruitment of Staff in Subordinate Courts is organize Clerk written test for District and Sessions Judges courts. So they all applicants will be select after category wise cut off marks decide. Below you may check estimated hc Haryana clerk exam 2019.
|Sr. No.
|Cutoffs Marks
|General / Unreserved Cat.
|S.C of Punjab
|Others S.C of Punjab
|BC / OBC of Punjab
|PHC of Punjab
|ESM (General) of Punjab
|ESM (SC) of Punjab
|ESM (BC/OBC) of Punjab
|Note : The about cut off is final list after typing test & written exam of 25 and 26 Feb 2017 examination for Clerk Recruitment (as Previous years cut off history). New cut off will be provide soon.
Download Punjab and Haryana HC Clerk Result 2019 Marks – Releasing Soon
Download Punjab and Haryana HC Cut-off Marks 2019 : Releasing Soon
Steps to check HC of Punjab & Haryana Clerk Results 2019
- Step : First of all enter to authorized portal https://sssc.gov.in.
- Step : On main page got download results of clerk examination 2019.
- Step : Open that one or enter roll number or little bit of internet info.
- Step : Login with valid candidature information.
- Step : Download Score Card.
- Step : A separate file PDF download for cut off / selection status.


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10 Jan, 2013
How do you decide who to follow on Twitter ?
Posted by andrea tomkins in: The business of blogging
I think this probably happens to everyone who has an account that reaches a certain milestone, but suddenly I'm getting a whole bunch more Twitter followers than ever before. I'm not one of those people who automatically follows everyone back. I spend about 5 seconds deciding whether I will follow someone or not. This is what runs through my mind:
- Has the person posted a photo of themselves? If it's blank I automatically assume they're a spammer. I also prefer photos of people, not logos, or cats, because I prefer to see a friendly face. Speaking of which, I have unfollowed people with gross and/or animated photos because I can't stand seeing them in my feed. So pick a good photo of yourself and run with it.
- What's the twitter handle? If it's something like SunnyDay23765 I assume it's a spammer and don't read on.
- Has the person filled out a bio? Some people don't bother, so I don't bother.
- How did they use their allotted 160 characters for their bio? The bio is make it or break it for me. And this is where I confess that I don't often follow people in insurance/car/real estate/pyramid-type sales. I don't want to open myself up to the hard sell. Sorry, but it's true. If you list yourself as a salesperson first above all other things I assume it's your priority and that you're just out to find clients. Be creative. Be yourself. Scratch that… be the Best Creative Version of yourself. And use good keywords too. If you love cupcakes and want to connect with other cupcake lovers, say so in your bio and they will find you.
- Has the person told me where they live? I am more likely to follow someone who lives in my home town. Filling in a location can also create some work/networking/interpersonal opportunities too, so if you are the kind of person who is looking to boost your Twitter followers, it might be a good idea to reveal where you are tweeting from.
- What has the person been tweeting lately? If it's funny, relevant, insightful, or interesting, I am more likely to follow. I am also more likely to follow if there is a good balance of "look at me" tweets and tweets about other people. I have unfollowed people who talk about themselves ad nauseum. Twitter is a conversation, and I don't want to follow people who just talk about themselves all the time. And don't be a jerk. Because if I see you being a jerk (to me, or to anyone one Twitter) I will never give you my business. In fact, I'll tell people to stay away from you because you're such a jerk. Here's a good exercise: open your twitter feed and look at your last 5-10 tweets. Do those tweets reflect the kind of person you are?
- I am lukewarm in regards to inspirational quotes from Gandhi, Einstein, Maya Angelou, etc. but I think opinions are mixed on this one.
- Has the person interacted with me? I may not follow the account at first, but if I sense that they're listening and interacting with me I am more likely to follow.
If you're on Twitter I'd love to hear how you decide who to follow.
p.s. I should probably mention that I'm @missfish on Twitter eh? :)