A novel efficient producer of human ω-amidase (Nit2) in Escherichia coli.

Nit2/ω-amidase catalyzes the hydrolysis of α-ketoglutaramate (KGM, the α-keto acid analogue of glutamine) to α-ketoglutarate and ammonia. The enzyme also catalyzes the amide hydrolysis of monoamides of 4- and 5-C-dicarboxylates, including α-ketosuccinamate (KSM, the α-keto acid analogue of asparagine) and succinamate (SM). Here we describe an inexpensive procedure for high-yield expression of human Nit2 (hNit2) in Escherichia coli and purification of the expressed protein. This work includes: 1) the design of a genetic construct (pQE-Nit22) obtained from the previously described construct (pQE-Nit2) by replacing rare codons within an 81 bp-long DNA fragment "preferred" by E. coli near the translation initiation site; 2) methods for producing and maintaining the pQE-Nit22 construct; 3) purification of recombinant hNit2; and 4) activity measurements of the purified enzyme with KGM and SM. Important features of the hNit2 gene within the pQE-Nit22 construct are: 1) optimized codon composition, 2) the presence of an N-terminus His6 tag immediately after the initiating codon ATG (Met) that permits efficient purification of the end-product on a Ni-NTA-agarose column. We anticipate that the availability of high yield hNit2/ω-amidase will be helpful in elucidating the normal and pathological roles of this enzyme and in the design of specific inhibitors.

Phenotyping of Potato Plants Using Morphological and Physiological Tools.

Potato (Solanum tuberosum L.) is one of the main non-grain agricultural crops and one of the main sources of food for humanity. Currently, growing potatoes requires new approaches and methods for cultivation and breeding. Phenotyping is one of the important tools for assessing the characteristics of a potato variety. In this work, 29 potato varieties of different ripeness groups were studied. Linear leaf dimensions, leaf mass area, number of stems, number of tubers per plant, average tuber weight, signs of virus infection, dry weight, pigment content, and number of stomata per unit leaf area were used as phenotyping tools. The strongest positive relationship was found between yield and bush area in the stage of full shoots (R = 0.77, p = 0.001), linear dimensions of a complex leaf (R = 0.44, p = 0.002; R = 0.40, p = 0.003), number of stems (R = 0.36, p = 0.05), and resistance to viruses X (R = 0.42, p = 0.03) and S (R = 0.43, p = 0.02). An inverse relationship was found between growth dynamics and yield (R = -0.29, p = 0.05). Thus, the use of morphological and physiological phenotyping tools in the field is informative for predicting key agricultural characteristics such as yield and/or stress resistance.

Detergent-Resistant Membranes in Chloroplasts and Mitochondria of the Halophyte Salicornia perennans under Salt Stress.

Halophytes represent important models for studying the key mechanisms of salt tolerance. One approach to the development of new knowledge of salt tolerance is to study the properties of detergent-resistant membranes (DRMs). In this work, the lipid profiles of DRMs of chloroplasts and mitochondria of euhalophyte Salicornia perennans Willd, before and after their exposure to shock concentrations of NaCl, have been investigated. We found that DRMs of chloroplasts are enriched in cerebrosides (CERs) and that sterols (STs) dominate the mass of mitochondrial DRMs. Also, it has been proven that (i) the impact of salinity provokes obvious growth in the content of CERs in DRMs of chloroplasts; (ii) the content of STs in DRMs of chloroplasts does not change under the influence of NaCl; (iii) salinity also causes some elevation in the content of monounsaturated and saturated fatty acids (FAs). Considering the fact that DRMs represent integral parts of both chloroplast and mitochondrial membranes, the authors have come to the conclusion that the cells of euhalophyte S. perennans, under the impact of salinity, presumes the choice (by the cell) of some specific composition of lipids and FAs in the membrane. This may be considered as a specific protection reaction of the plant cell against salinity.

Effect of Salinity on Leaf Functional Traits and Chloroplast Lipids Composition in Two C3 and C4 Chenopodiaceae Halophytes.

Salt stress is one of the most common abiotic kinds of stress. Understanding the key mechanisms of salt tolerance in plants involves the study of halophytes. The effect of salinity was studied in two halophytic annuals of Chenopodiaceae Salicornia perennans Willd. and Climacoptera crassa (Bied.) Botsch. These species are plants with C3 and C4-metabolism, respectively. We performed a comprehensive analysis of the photosynthetic apparatus of these halophyte species at different levels of integration. The C3 species S. perennans showed larger variation in leaf functional traits-both at the level of cell morphology and membrane system (chloroplast envelope and thylakoid). S. perennans also had larger photosynthetic cells, by 10-15 times, and more effective mechanisms of osmoregulation and protecting cells against the toxic effect of Na+. Salinity caused changes in photosynthetic tissues of C. crassa such as an increase of the mesophyll cell surface, the expansion of the interface area between mesophyll and bundle sheath cells, and an increase of the volume of the latter. These functional changes compensated for scarce CO2 supply when salinity increased. Overall, we concluded that these C3 and C4 Chenopodiaceae species demonstrated different responses to salinity, both at the cellular and subcellular levels.

Balance of Δ5-and Δ7-sterols and stanols in halophytes in connection with salinity tolerance.

Sterols (STs) have a key role in regulating the fluidity and permeability of membranes in plants (phytosterols) that have wide structural diversity. We studied the effect of structural STs diversity on salt tolerance in halophytes. Specifically, we used gas chromatography-mass spectrometry (GC-MS), including two-dimensional gas chromatography-mass spectrometry (GCxGC-MS), to assess the STs composition in leaves of 21 species of wild-growing halophytes from four families (Asteraceae, Chenopodiaceae, Plumbaginaceae, Tamaricaceae) and three ecological groups (Euhalophytes (Eu), recretophytes (Re), salt excluders (Ex)). Fifteen molecular species of STs from three main groups, Δ5-, Δ7-and Δ0- STs (stanols), were detected. Plants of the genus Artemisia were characterized by a high content of stigmasterol (30-49% of the total STs), while ß-sitosterol was the major compound in two Limonium spp., where it comprised 84-92% of the total STs. Species of Chenopodiaceae were able to accumulate both Δ5-and Δ7-STs and stanols. The content of the predominant Δ5-STs decreased in the order Ex → Re → Eu. Molecular species with a saturated steroid nucleus were identified in Eu and Re, suggesting their special salt-accumulating and salt-releasing functions. The structural analogues of stigmasterol, having a double bond C-22, were stigmasta-7,22-dien-3ß-ol (spinasterol) and stigmast-22-en-3ß-ol (Δ7--sitosterol). The ratio of Δ5-stigmasterol/Δ5-ß-sitosterol increased in Ex plants, and spinasterol/Δ7--sitosterol and 22-stigmastenol/sitostanol increased in Eu plants. These data support the well-known role of stigmasterol and its isomers in plant responses to abiotic and biotic factors. The variability in STs types and their ratios suggested some involvement of the sterol membrane components in plant adaptation to growth conditions. The balance of Δ5-, Δ7-and stanols, as well as the accumulation of molecular analogues of stigmasterol, was suggested to be associated with salt tolerance of the plant species in this investigation.

Heterotrophic Microbiota from the Oligotrophic Waters of Lake Vostok, Antarctica.

Lake Vostok is the deepest lake of Antarctica but has poor accessibility for study due to a thick glacial cover, however, water samples of this lake have become available for study just recently. Previously, only the microbiome of the ice cover samples was characterized. Here we report results of bacteriological seeding with subsequent identification of the heterotrophic microorganisms (bacteria and micellar fungi) present by 16S rDNA sequencing as well as results of a direct molecular study of the water microbiome. Surprisingly, the data obtained gave evidence of a predominant occurrence of common chemoorganotrophs that were rather psychrotolerant than psychrophilic. We isolated and described strains belonging to eight heterotrophic microbial species able to grow in a rich medium: six bacterial strains belonging to the species Microbacterium testaceum and Microbacterium trichothecenolyticum, Brevundimonas diminuta, Sphingomonas oligophenolica, Sphingomonas sp. and Sphingobium limneticum; and two fungal strains belonging to Dendryphion sp. and Cladosporium fusiforme. Direct study of 16S rDNA purified water samples confirmed the predominance of the Brevundimonas, Microbacterium, Bradyrhizobium, and Bacillus (Bacillus cereus) genera.

Effect of NaCl, copper and cadmium ions on halophytes with different types of salt resistance: accumulation, physiological and biochemical reactions.

The capacities of the euhalophyte SaLi-Cornia perennans Willd. and glycohalophyte Artemisia santonica L. to accumulate NaCl, Cu, and Cd, as well as their physiological and biochemical responses to these compounds, was investigated. Seeds were germinated in distilled water for 1-3 days and then sown in containers with sand. Plants were watered with Robinson's nutrient solution. After 3 months, plants were divided into two groups: experimental and control. In the experimental group, soil was treated with 1M NaCl, 10 mM Cu(NO3)2, and 10 mM Cd(NO3)2 for 24 h. The exposure to high concentration of NaCl in the experiment did not affect the baseline level of Na, which was twice as high in S. perennans as in A. santonica. Plant exposure to Cu and Cd caused their accumulation in the aboveground parts of both species. The accumulation capacity of the euhalophyte was many times higher than that of the glycohalophyte. We analysed functional parameters of leaves by measuring photosynthetic pigments, structural parameters of membranes by assessing the lipid profile, and the balance of pro/antioxidant processes. Using data on changes in several biochemical parameters, the sensitivity of the two different halophytes to metal ions was as follows: for S. perennans - Cu > Na > Cd; for A. santonica - Na > Cu > Cd. Our findings suggest that S. perennans can be used for heavy metal extraction from soil in phytoremediation, whereas A. santonica will be more effective for greening of polluted territories.

Indigenous women's reproductive health in the Arctic zone of Western Siberia: challenges and solutions.

In the Russian Arctic, alarming trends (shortage of nomadic Indigenous women, high reproductive loss, child mortality rates) indicate long-term changes towards demographic decline. This study aimed at comparing some indicators of the reproductive health (childbirth rates, number of pregnancies, pregnancy loss) of Indigenous and non-Indigenous women in the exploration of cultural and social factors influencing reproductive behaviour. A multidisciplinary approach draws on methods of medicine, sociology and health economics. It includes data of the women's reproductive health collected from surveys of 879 women (of whom 627 were Indigenous) during expeditions to the settlements and the tundra of the Yamal-Nenets Autonomous Okrug in 2013-2019. In the tundra, 66.7% of registered Indigenous women's pregnancies resulted in childbirth, 7.8% in induced abortions, 25.5% in spontaneous miscarriage. More than three children were delivered by 59.1% of Indigenous women. Most Indigenous families suffered from high pregnancy loss. Marriage between close relatives was 27.0%. Child mortality equalled 24.4%, three times higher than in the population of the settlements and eight times higher than in families immigrating from other regions. The survival of Indigenous peoples significantly depends on women's reproductive health and sufficient medical service that requires targeted policy.

Changing diets and traditional lifestyle of Siberian Arctic Indigenous Peoples and effects on health and well-being.

The diet of Indigenous Peoples of North-Western Siberia is characterized by a significant proportion of traditional foods. Eating local products provides a ready-made set of macro- and microelements necessary for life in the challenging conditions of the Arctic. Currently, high consumption of traditional foods is typical in the season of fishing or reindeer slaughter, while out of season the consumption of easily digestible carbohydrates increases. Due to climate change, seasonal fishing and traditional migration routes are disrupted and, therefore, the consumption of traditional foods is decreasing. During 5 years of expeditions, we performed a cross-sectional screening of 985 Indigenous People from three districts of the Yamal-Nenets Autonomous Okrug. We analyzed the seasonality of acquisition and consumption of local reindeer and fish products and studied the traditional food storage among 90 Indigenous fishermen and hunters, who exploit cryostorage. As diet affects health, we studied the benefits of local food. The duration of the consumption season is decreasing and therefore the amount of consumption of local food is decreasing as well. This has adverse effects on health with increasing hypertension dissemination. The creation of stocks of fish and reindeer meat in villages and their year-round sale to the population is a necessary step for ensuring the food security and health of the inhabitants in the region.

Food Sovereignty of the Indigenous Peoples in the Arctic Zone of Western Siberia: Response to COVID-19 Pandemic.

This article presents the challenges facing reindeer herding as being both a profitable business and part of the traditional culture of the nomadic Indigenous peoples in the Arctic zone of Western Siberia which addresses substantial needs of the local population. Reindeer herding products are used as traditional nutrition, and as effective preventive means and remedies for adapting to the cold and geomagnetic activity in the High North. Export trends of traditional reindeer products have decreased local Indigenous peoples' access to venison and had a negative impact on their health. Due to the COVID-19 pandemic, it is especially urgent for the Indigenous peoples to have sufficient access to traditional food and be involved in policy decision-making to maintain this traditional business. We aim to analyze the dependencies of Indigenous peoples on the reindeer produce-exporting "food value chain" and explore how (1) the independence of reindeer herders could be increased in these export chains and (2) how provision of their products to local communities could be secured. The study takes a multidisciplinary approach based on policy and socioeconomic analyses with input from medical research. Primary sources include data collected from interviews and surveys of Indigenous peoples during expeditions to the Nyda settlement, the Nydinskaya tundra, the Tazovsky settlement, the Tazovskaya tundra, the Nakhodka tundra, the Gyda and Gydansky settlements, the Yavai-Salinskaya tundra, the Seyakha settlement, the Seyakhinskaya and Tambeyskaya tundras located along the southern coast of the Ob Bay, the northeast coast of the Yamal Peninsula, the Tazovsky and Gydansky Peninsulas, and the Shuryshkarsky district. Data were collected during the summers and winters of 2014-2020.

Structural and molecular strategy of photosynthetic apparatus organisation of wild flora halophytes.

Structural and molecular parameters of photosynthetic apparatus in plants with different strategies for the accumulation of salts were investigated. CO2 gas exchange rate, content of pigments, mesostructure, chloroplast ultrastructure and the biochemical composition of the membrane structural components in leaves were measured. The objects of the study were euhalophytes (Salicornia perennans, Suaeda salsa, Halocnemum strobilaceum), crynohalophyte (Limonium gmelinii), glycohalophyte (Artemisia santonica). Euhalophytes S. perennans and S. salsa belong to the plants of the halosucculent type, three other species represent the xerophilic type. The highest photosynthetic activity estimated by the average parameters of CO2 gas exchange rate in the leaves was observed in S. perennans plants. Plants of the xerophyte type including both H. strobilaceum euhalophyte and cryno- and glycohalophytes are described by lower values of these characteristics. Larger cells with a great number of chloroplasts and a high content of membrane glycerolipids and unsaturated C18:3 fatty acid, but with smaller pigment and light-harvesting complexes size characterise the features of euhalophytes with a succulent leaf type. Thus, features of the mesostructure, ultrastructure, and supramolecular interactions of the halophyte PA were closely related to the functional parameters of gas exchange, and were characterised by the strategy of species in relation to the accumulation of salts, the life form of plants, and the attitude to the method of water regulation.

Cognition, emotion, and arithmetic in primary school: A cross-cultural investigation.

The study investigated cross-cultural differences in variability and average performance in arithmetic, mathematical reasoning, symbolic and non-symbolic magnitude processing, intelligence, spatial ability, and mathematical anxiety in 890 6- to 9-year-old children from the United Kingdom, Russia, and China. Cross-cultural differences explained 28% of the variance in arithmetic and 17.3% of the variance in mathematical reasoning, with Chinese children outperforming the other two groups. No cross-cultural differences were observed for spatial ability and mathematical anxiety. In all samples, symbolic magnitude processing and mathematical reasoning were independently related to early arithmetic. Other factors, such as non-symbolic magnitude processing, mental rotation, intelligence, and mathematical anxiety, produced differential patterns across the populations. The results are discussed in relation to potential influences of parental practice, school readiness, and linguistic factors on individual differences in early mathematics. Statement of contribution What is already known on this subject? Cross-cultural differences in mathematical ability are present in preschool children. Similar mechanisms of mathematical development operate in preschool children from the United Kingdom, Russia, and China. Tasks that require understanding of numbers are best predictors of arithmetic in preschool children. What does this study add? Cross-cultural differences in mathematical ability become greater with age/years of formal education. Similar mechanisms of mathematical development operate in early primary school children from the United Kingdom, Russia, and China. Symbolic number magnitude and mathematical reasoning are the main predictors of arithmetic in all three populations.

Efficacy and safety of rituximab combined with ESHAP chemotherapy for the treatment of relapsed/refractory aggressive B-cell non-Hodgkin lymphoma.

BACKGROUND: We evaluated the efficacy and safety of adding rituximab to nonanthracycline ESHAP (etoposide/methylprednisolone/cytarabine/cisplatin) chemotherapy for relapsed/refractory aggressive non-Hodgkin lymphoma (NHL). PATIENTS AND METHODS: Patients with intermediate- or high-grade NHL were to receive 6 rituximab doses and 6 ESHAP cycles. Rituximab 375 mg/m(2) was administered 1 week and 1 day before cycle 1 of standard ESHAP (etoposide 40 mg/m(2) on days 1-4; methylprednisolone 500 mg/m(2) on days 1-5; cytarabine 200 mg/m(2) on day 5; and cisplatin 25 mg/m(2) on days 1-4). Rituximab was repeated before the third and fifth 21-day ESHAP cycles (on days 48 and 90 of protocol, respectively), followed by 2 additional rituximab doses after cycle 6 (on days 134 and 141 of protocol). Use of growth factors was permitted. Thirteen patients were enrolled (median age, 56 years); all had previously treated NHL, 12 (92%) had diffuse large B-cell lymphoma, 10 (77%) had stage III/IV disease, and 2 (15%) had chemotherapy-refractory disease. RESULTS: The most common grade 3/4 toxicities were neutropenia and thrombocytopenia, with 3 cases of febrile neutropenia. Seven patients exhibited complete response (CR) and 3 had partial response, for an objective response rate of 77%. Median duration of response for all responders was 14 months (range, 2-51 months). Among 6 patients completing all 6 cycles, 4 (67%) had a CR, 1 had a partial response, and 1 had progressive disease. Three of the 4 CRs have remained for a median of 48 months (range, 46-51 months). CONCLUSION: Rituximab plus ESHAP led to durable responses with acceptable toxicity in patients with relapsed/refractory aggressive NHL, most of whom had advanced disease.

Photosynthetic parameters and redox homeostasis of Artemisia santonica L. under conditions of Elton region.

Structural and functional parameters and redox homeostasis in leaves of Artemisia santonica L. under environment conditions of Elton lake (the southeast region of the European part of Russia) were measured. The highest photosynthetic apparatus (PA) activity in A. santonica leaves on CO2 gas exchange as well as the highest content of green pigments was observed in the morning. Maximum share of violaxanthin cycle key pigments - zeaxanthin (Zx) and antheraxanthin (Ax) was observed in the afternoon and decreased in the evening. Lipids/chlorophyll (Chl) ratio increased in the evening due to the decrease in Chl concentration, and content of linolenic acid (С18:3n3) was decreased in the middle of the day. The content of TBA-reacting products increased 1.4-fold in the middle of the day, and decreased approximately 2-fold in the evening. The decrease of the activity was observed in diurnal dynamics of superoxide dismutase (SOD) and polyphenol oxidase (PPO). Increased accumulation of phenols and flavonoids, as well as free amino acids (FAA) in A. santonica leaves was observed in the middle of the day. Thus, the ability of A. santonica plants to resist the soil salinization, high levels of solar illumination and temperature consists of a number of protectively-adaptive reactions of metabolic and photosynthetic control.

Seasonal dynamic of morpho-physiological properties and the lipid composition of Plantago media (Plantaginaceae) in the Middle Volga region.

The changes in morpho-physiological properties and lipid composition have been studied in the leaves of the plant Plantago media collected from two different places in the Middle Volga region during the summer of 2010. The plants gathered from the first plot (P1 plants) grew on plain ground in the midst of typical meadow-steppe perennial plants. The plants of the second group (P2 plants) grew on a flat slope of the South-West exposition, in the grass community. The leaves of the plants Р1 had lower specific area densities but larger areas and masses; they accumulated more levels lipid peroxide products. The changes in lipid compositions depended on the growth phase and habitats. Correlations between morpho-physiological parameters and certain lipids have been established. The amounts of galactolipids (GL) have been shown to correlate with the leaf areas. When the leaf areas were reduced, a ratio between phosphatidylcholines (PC) and phosphatidylethanolamines (PE) decreased. The result of our study showed that gradual changes of morphometrical parameters were accompanied by the alterations in biomass structure and modifications in lipids and fatty acids (FA).

The Impact of a Line Probe Assay Based Diagnostic Algorithm on Time to Treatment Initiation and Treatment Outcomes for Multidrug Resistant TB Patients in Arkhangelsk Region, Russia.

BACKGROUND: In the Arkhangelsk region of Northern Russia, multidrug-resistant (MDR) tuberculosis (TB) rates in new cases are amongst the highest in the world. In 2014, MDR-TB rates reached 31.7% among new cases and 56.9% among retreatment cases. The development of new diagnostic tools allows for faster detection of both TB and MDR-TB and should lead to reduced transmission by earlier initiation of anti-TB therapy. STUDY AIM: The PROVE-IT (Policy Relevant Outcomes from Validating Evidence on Impact) Russia study aimed to assess the impact of the implementation of line probe assay (LPA) as part of an LPA-based diagnostic algorithm for patients with presumptive MDR-TB focusing on time to treatment initiation with time from first-care seeking visit to the initiation of MDR-TB treatment rather than diagnostic accuracy as the primary outcome, and to assess treatment outcomes. We hypothesized that the implementation of LPA would result in faster time to treatment initiation and better treatment outcomes. METHODS: A culture-based diagnostic algorithm used prior to LPA implementation was compared to an LPA-based algorithm that replaced BacTAlert and Löwenstein Jensen (LJ) for drug sensitivity testing. A total of 295 MDR-TB patients were included in the study, 163 diagnosed with the culture-based algorithm, 132 with the LPA-based algorithm. RESULTS: Among smear positive patients, the implementation of the LPA-based algorithm was associated with a median decrease in time to MDR-TB treatment initiation of 50 and 66 days compared to the culture-based algorithm (BacTAlert and LJ respectively, p<0.001). In smear negative patients, the LPA-based algorithm was associated with a median decrease in time to MDR-TB treatment initiation of 78 days when compared to the culture-based algorithm (LJ, p<0.001). However, several weeks were still needed for treatment initiation in LPA-based algorithm, 24 days in smear positive, and 62 days in smear negative patients. Overall treatment outcomes were better in LPA-based algorithm compared to culture-based algorithm (p = 0.003). Treatment success rates at 20 months of treatment were higher in patients diagnosed with the LPA-based algorithm (65.2%) as compared to those diagnosed with the culture-based algorithm (44.8%). Mortality was also lower in the LPA-based algorithm group (7.6%) compared to the culture-based algorithm group (15.9%). There was no statistically significant difference in smear and culture conversion rates between the two algorithms. CONCLUSION: The results of the study suggest that the introduction of LPA leads to faster time to MDR diagnosis and earlier treatment initiation as well as better treatment outcomes for patients with MDR-TB. These findings also highlight the need for further improvements within the health system to reduce both patient and diagnostic delays to truly optimize the impact of new, rapid diagnostics.

Membrane-forming lipids of wild halophytes growing under the conditions of Prieltonie of South Russia.

The composition of membrane-forming lipids has been examined for 10 wild halophyte species growing in southern Russian on alkaline soil. The plants belong to seven taxa of family rank: by their life form, which are semi-shrubs, herbaceous annuals, and perennial plants; their salt tolerance, which are classified as the euhalophytes, crynohalophytes, and glycohalophytes; and by their sensitivity to water, classifications of mesoxerophytes and xeromesophytes. Parallels have been found between the lipid composition and the ecological status of the plants. It has also been revealed that the similarity in the glyco- and phospholipid composition of different plant groups relates to the water factor and the type of salt accumulation, respectively. The fatty acid compositions of the examined plants is determined at the species level.

Tn5044-conferred mercury resistance depends on temperature: the complexity of the character of thermosensitivity.

Escherichia coli K12 containing the transposon Tn5044 mer operon (merR, T, P, C, and A genes) is resistant to mercuric chloride at 30 degrees C but sensitive to this compound at 37-41.5 degrees C. We have studied the mechanism underlying the temperature-sensitive nature of this mercury resistance phenotype, and found that the expression of the Tn5044 merA gene coding for mercuric reductase (MerA) is severely inhibited at non-permissive temperatures. Additionally, MerA showed a considerably reduced functional activity in vivo at non-permissive temperatures. However, the temperature-sensitive character of the functioning of this enzyme in cell extracts, where it interacted with one of the low-molecular weight SH compounds rather than with the transport protein MerT (as is the case in vivo), was not apparent. These data suggest that the temperature-sensitive mercury resistance phenotype should stay under control at two stages: when the merA gene is expressed and when its product interacts with MerT to accept the mercuric ion.