#trisaccharides

[cornley-amateur-drama-society]

CONTEXT: Max's side of this peace is fragile AU. He is talking to Lucy about the peace treaty offered by Beawynn.

Max let out a sigh, leaning heavily against the wall of the Castle Bennegrove, the siblings having decided to go out on a small hunt with several cousins later that afternoon.  He looked around for any wandering ears, before answering her question.  “He's set to tell the Court at the end of the week,” he started, rubbing absently at his wrist.  “I'm to be married.”

CONTEXT: The Techie's part of the trisaccharides trilogy!

“I can answer that,” Blessing said cheerfully, setting down a finished card in light blue with white ribbon decorating the front in little flowers.  “Rob decided that the CYDS Kids were going to have a Valentine’s Do after class got out earlier.”

this week's word is...

Find the word in any WIP and share the sentence containing it. Reply, reblog, stick it in the tags, tag us in a new post, or keep it private. All fandoms, all ships, all writers welcome.

[delifoodsmanufacturer]

What Is Isomalto-oligosaccharide Syrup(IMO Syrup)?

In recent years, sugar control, fat reduction, and healthy living have become important ingredient trends in baked pastries. Sucrose-free and natural sugar substitutes have gradually attracted attention, and consumers are pursuing healthier sweetness. As a new type of functional sweetener, Isomalto-oligosaccharide syrup is widely used in medical, functional food and food additives industries due to its unique health properties. It is called a new type of biological glycogen in the 21st century. Isomalto-oligosaccharide syrup is a functional oligosaccharide made from high-quality starch (corn or cassava) by enzymatic hydrolysis and refining. It has the characteristics of low sweetness, low calories, good moisture retention, and anti-caries, and is widely used in food, beverages, health products and other fields. Currently, two main types of isomaltose products are on the market: IMO-50 and IMO-90. The content of oligosaccharides in IMO-50 must account for more than 50% of the total dry matter, of which isomaltose, panose, and isomaltotriose account for more than 35% of the total dry matter; IMO-90 requires that the content of IMO is more than 90% and the content of trisaccharides is more than 45%. Wuhu Deli Food Co., Ltd., as a leading syrup manufacturer in China, has been committed to providing consumers with safe, healthy and high-quality products. We have been deeply involved in the field of oligosaccharides for many years, with advanced production equipment, professional technical teams and a complete quality management system. Product characteristics of oligosaccharides Low sweetness -Oligosaccharides are about 40%-50% of sucrose, with a refreshing taste and will not give people a feeling of being too sweet. This makes it more widely used in food and beverages, which can meet the taste needs of different consumers. Low calories- Compared with sucrose, oligosaccharides have lower calories, containing only about 2 kcal per gram. This is an ideal choice for consumers who are concerned about health and weight management. At the same time, the low-calorie characteristics also make oligosaccharide syrup more and more popular in low-sugar and low-fat foods. Good moisture retention-oligosaccharide syrup has good moisture retention, which can keep the moisture of food and beverages and extend their shelf life. In baked goods, oligosaccharide syrup can make bread, cakes and other products softer, moister and taste better. Anti-caries-oligosaccharide syrup cannot be fermented and utilized by bacteria in the mouth, so it has an anti-caries effect. This is a very important advantage for children and teenagers, which can effectively prevent the occurrence of caries. The application range of Isomalto-oligosaccharide syrup is very wide, mainly including the following aspects: Dairy products: yogurt, ice cream, milk powder, etc. Beverages: functional drinks, carbonated drinks, juices, etc. Baked foods: bread, biscuits, cakes, etc. Candy: soft candy, hard candy, etc. Condiments: sauces, seasoning powders, etc. Health food: probiotic food, dietary supplements, etc. At present, our syrup products are exported to Europe, America, Southeast Asia and other countries. Deli foods has been established for over 20 years and has always adhered to quality-oriented standards, providing customers with high-quality syrup products. As a food factory, it can effectively control production costs. Therefore, we can provide customers with reasonable prices, allowing customers to gain an advantage in market competition. We will provide each customer with high-quality products, low prices, fast delivery and professional services for win-win cooperation. If you need other syrups, such as rice syrup, fructose syrup, glucose syrup, etc., Welcome to contact us and request free samples~ Read the full article

[aayushi1806]

What Is Malt? And Malt Processing?

What Is Malt? And Malt Processing?

Malt is germinated cereal grain that has been dried in a process known as “malting”.  Malted barley, or ‘Malt’ as it is most commonly known, is a wonderful package of starch, enzymes, protein, vitamins, and minerals plus many other minor constituents that provide the brewer and distiller with their main raw material.

The grains are made to germinate by soaking in water and are then halted from germinating further by drying with hot air. Malting grains develop the enzymes required for modifying the grain’s starches into various types of sugar, including monosaccharide glucose, disaccharide maltose, trisaccharide malt triose, and higher sugars called maltodextrins.

It also develops other enzymes, such as proteases, which break down the proteins in the grain into forms that can be used by yeast. Depending on when the malting process is stopped, one gets a preferred starch to enzyme ratio and partly converted starch into fermentable sugars. Malt also contains small amounts of other sugars, such as sucrose and fructose, which are not products of starch modification but were already in the grain. Further conversion to fermentable sugars is achieved during the mashing process.

Malted grain is used to make beer, whiskey, malted milkshakes, malt vinegar, confections such as Maltesers and Whoppers, flavoured drinks such as Horlicks, Ovaltine, and Milo, and some baked goods, such as malt loaf, bagels, and rich tea biscuits. Malted grain that has been ground into a coarse meal is known as “sweet meal”. Various cereals are malted, though barley is the most common. A high-protein form of malted barley is often a label-listed ingredient in blended flours typically used in the manufacture of yeast bread and other baked goods.

60-65% of the weight of malt is un-degraded starch and malt contains all the key enzymes for starch degradation during the mashing stage of both the brewing and distilling process. These enzymes produce fermentable sugars to supplement the other key nutrients for yeast growth that malt provides. These include amino acids, vitamins, and minerals.

The malting process consists of 4 stages steeping, germination, kilning and roasting.

1. Steeping

The basic malting process, although more of an exact science today than when man first dipped baskets of grain into open wells in Mesopotamia 5,000 years ago to prepare it for brewing, remains a three-step process: steeping, germination, and drying.

During steeping water is absorbed by the raw barley kernel and germination begins. Steeping starts with raw barley that has been sorted and cleaned, then transferred into steep tanks and covered with water. For the next 40-48 hours, the raw barley alternates between submerged and drained until it increases in moisture content from about 12% to about 44%. The absorbed water activates naturally existing enzymes and stimulates the embryo to develop new enzymes. The enzymes break down the protein and carbohydrate matrix that encloses starch granules in the endosperm, opening up the seed’s starch reserves, and newly developed hormones initiate growth of the acrospires (sprout).

Steeping is complete when the barley has reached a sufficient moisture level to allow a uniform breakdown of the starches and proteins. One visual indicator that the maltster uses to determine the completion of steeping is to count the percentage of kernels that show “chit”. Raw barley that has been properly steeped is referred to as “chitted” barley”, the “chit” being the start of the rootlets that are now visibly emerging from the embryo of the kernel.

2. Germination

Germination is the ‘control’ phase of malting. It continues for a further 4-5 days depending on the product type being made. The germinating grain bed is kept at a temperature and oxygenated by providing a constant flow of humidified air through the bed at specific temperatures. The grain is turned regularly to prevent rootlets matting and to maintain a loosely packed grain bed. The maltster manipulates the germination conditions to vary the type of malt being manufactured.

3.Kilning

Kilning, the third phase of malting, dries the grain down to 3-5% moisture and arrests germination. Large volumes of hot air are blown through the grain bed. By varying air flows and kiln temperatures, malts of different colours can be produced with varying flavour profiles. At the end of kilning the malt is cooled and the tiny rootlets removed before analysis and storage. The final malt is analysed extensively according to malt type and customer profile. The malt may be dispatched in bags, in containers or in bulk.

4. Roasting

Roasting is done in 4 distinct stages: steeping, germinating, roasting and cooling. At GWM Malt, grain spends 34-46 hours in steep tanks where we aim for target moisture of 42-44%. The grain is transferred to germination which lasts for around 4 weeks. This is a semi-continuous moving batch germination process. Once germination is complete, the green malt is then transferred to the roasting drum.

The roasting takes place in two roasting drums. The average roasting time is 2 ½ – 3 hours with air on temperatures of up to 460˚C. Our roasters take a batch size of 2.4 – 3.5 tonnes. The roasted malt is then transferred to the cooler and spends 35 – 60 minutes there in order to drop the temperature to <15˚C and fix the colour and flavour compounds. The malt is analysed before storage and thereafter awaits dispatch to our customers.

Benefits Of Malt:

1. Rich In B Vitamins

As a rich source of B vitamins, malt extract may increase the B-vitamin content of the beverages it’s used in — including thiamine, riboflavin, niacin, folate, and vitamin B-6. The amount may vary depending on the malt beverage you’re drinking, however. B vitamins are necessary for metabolising the carbohydrates, protein, and fat in food into energy. They also help regulate appetite, promote good vision and keep your skin healthy.

2. Source Of Essential Amino Acids

Malt extract is a source of essential amino acids, which your body needs to make the proteins. Although some malt extract beverages are not a significant source of protein, they may help provide a small amount of these essential nutrients, boosting your intake.

3. Good For Your Bones

Good nutrition is important for bone health. Some malt extract beverages may not be a significant source of the nutrients your bones need for good health, but they can help boost your intake. In addition to calcium, these drinks may also contain phosphorus and magnesium, also important minerals that help keep your bones healthy and strong. All three minerals make up the primary structure of your bones, while magnesium is also needed to regulate the hormones responsible for mineral metabolism.

4. Things To Consider

While malt extract beverages offer some nutritional benefits, they may not be a significant source of many of these health-promoting nutrients. It’s OK to include such beverages in your regimen, but they should be consumed as part of an overall healthy diet so your body gets all the nutrients it needs from a variety of sources.

[cornley-amateur-drama-society]

it's only been a day but how did i lose all of my braincells for these two parts i *standing emoji*

alright alright alright! time to work on the cyds kids' part and the techie's part!

[moleculardepot]

Blood Group H type II Trisaccharide Biotinylated

Blood Group H type II Trisaccharide Biotinylated Catalog number: B2012967 Lot number: Batch Dependent Expiration Date: Batch dependent Amount: 1 mg Molecular Weight or Concentration: 926.0 g/mol Supplied as: Lyophilized Powder Applications: molecular tool for various biochemical applications Storage: -20°C Keywords: Fuca1-2Galb1-4GlcNAc-b-1-O(CH2)3NHCO(CH2)5NH-biotin Grade: Biotechnology grade.…

[0ctogus]

Another fun fact i was taught is that legumes and some other veggies contain raffinose, which is a trisaccharide, so not a veeery complex carbohydrate. Nonetheless humans lack the enzymes to digest it, so when it reaches the colon, it gets fermented and that's why legumes can cause gas!

FUN FACT TIME !!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!!

So, when you eat carbohydrates that cannot easily be broken down (by hydrolysis*) into glucose molecules, they don't get released into glucose molecules in your small intestine as would, for example, simpler sugars like sucrose that is only two monosaccharides (single sugars) long.

This means that these carbohydrates reach your large intestine not fully broken down. So, they tend to be fermented by gut bacteria instead. This has the lovely side effect of making one fart, as well as the genuinely lovely side effect of causing gut bacteria to (A) generally be healthier and (B) produce butanoic acid salts (known as butyrates), which are really important to regulate metabolism, and also produces SCFAs and idk what they do but Wikipedia makes it sound like they're good.

This is also why fibres (which is a broad term including things like cellulose*, so non-starch carbohydrates that aren't as easy to hydrolyse) are good for you - because they are a little bit tougher to digest, so they reach your large intestine where they are fermented by gut bacteria rather than simply instantly getting broken down into glucose the second they encounter a teeny bit of amylase.

But it gets even more interesting than that!

Starches that cannot be easily broken down are called Resistant Starches, right? Resistant starches include amylose. Amylose is a long straight chain of glucose molecules, which contrasts with the other type of starch, amylopectin, which has branches.

Because of its branching, amylopectin has a high surface area to volume ratio, so it is easier to digest. Amylose has a lower surface area to volume ratio so it is roughage and is trickier to digest, so it reaches the large intestine.

Also, in plant cells, starch is often stored in granules. What do we do when we cook food? The heat causes granules to expand, start leaking, or even burst completely, thus making our food easier to digest. It is harder to digest if you have to eat through the granule first before you can even START to break down the polymers. Cooking means that often times, the starch is Literally Right There, so it makes the food much easier to digest.

Anyway, stuff that is Really easy to digest, you get the sugar all at once, so it goes into storage or you get super energetic but it doesn't give you a good lasting amount of energy like slower-releasing starches do.

This all explains... like everything that people say about how you need to eat healthy. (Except for the stupid things like that you need to cut carbs.) It explains why fibre helps digestion, why more complex carbohydrates are often healthier than simple ones like sucrose, WHY WE COOK FOOD!!!!!!!!!!!!!!!!! IT'S SO COOL

Explanations for those who are confused by terminology under the cut:

*cellulose is found in cell walls and is a carbohydrate that is really tough since it forms a lattice shape. It is the stuff that makes wood so strong, and also forms part of lignin AKA tree bark.

*Basically, carbohydrates are made of single sugars that bond together by condensation. Condensation reactions are when on the end of two monomers, there is an OH group and an OH group, and then one of the OHs gets removed and another H+ off of the other OH is removed. This means both monomers are then sharing the one oxygen left, and there is a water molecule produced hence the name. Hydrolysis is the inverse of this - when a water molecule is split into OH- and H+ and then it breaks apart a polymer.

[maltextract123]

BENEFITS OF MALT EXTRACT at Mahalaxmi Malt Extract

What is Malt?

Malt- A grain when soaked in water and then dried in a hot oven produces a product known as Malt. So basically barley malt is produced through a process known as Malting in which steeping, germinating and drying of grain is done to convert it into malt.

It develops enzymes which are required for converting the starch present in grain into different types of sugars such as trisaccharide maltotriose, disaccharide, monosaccharide glucose, maltose,, and higher sugars called maltodextrins. Other than these, it also develops enzymes, such as proteases, that break down the proteins in the grain into forms that can be used by yeast.

What is Malt Extract?

Malt extract refers to the amount of soluble material that can be extracted from the malt during mashing

Why Mahalaxmi Malt Extract?

During various surveys, it's been asked from brewers to share their advice and experience for first time brewing. All of them suggested that the most time consuming step and the one that the new brewers should skip is the step of mashing.

As a brewer, you must be aware that malted grains are the most important ingredient in brewing and brewing involves the manufacture of beer from grains. Mashing is the term given to the start of the brewing process, where to form a porridge-like mixture crushed grains are mixed with water and the resulting product is called the “mash.

But wait! This sounds a bit complex right? No worries…. we Mahalaxmi Malt Extract are here to help you skip the step of mashing, by providing the best quality malt extract for brewing.

When you use Mahalaxmi Malt Extract, you can directly skip the step mashing. Both forms of malt extract are provided here- dry and liquid malt extract and you can use either one or both as per your recipe’s requirement.

Benefits:

Using Malt extract not only saves time but also provides you enormous benefits such as:

Saves Time – Only by skipping the mashing process, one can reduce the time taken for brewing substantially

Saves Space & Money – Since you don’t need the equipment used in mashing you don’t have to invest in large equipment like a mash tun, boiling kettle, etc.

Improves Consistency – Easily hit the target gravity every time you brew a new batch.

Improves Efficiency – One of the biggest challenges and roadblock for beginner brewers is keeping the alcohol levels in check. And with the help of malt extract, you have more control over the process.

Simplifies the Brewing Process – Especially for beginners

For more information about: Malt Extract Please visit at https://www.mahalaxmimaltextract.com/

[autodaemonium]

hərʌwaʊaɪiuvnɛziɪəbynw

Pronounced: huhruwowaiiuvnayziiuhbynw.

Pantheon of: subjectivism, ability, painfulness, lawfulness, chemical element, south, immobility, inheritance.

Entities

Aɪæssætsəfrbənrʃnkpəə

Pronounced: aiassatsuhfrbuhnrshnkpuhuh Lawfulness: legality. Inheritance: upbringing. Immobility: inertness. Chemical Element: ununpentium. Painfulness: sharpness. Ability: form. Legends: cat's cradle, dirty war, exacerbation, excursion. Prophecies: smuggling, exhortation, greyhound racing, least effort. Relations: ʃtnwgwbvhkstugiʌwəhɪ (transcriptase).

Gsəhɛpsðntvʌwlɪrɪlmy

Pronounced: gsuhhaypsthntvuwlirilmy Lawfulness: legitimacy. Inheritance: upbringing. Immobility: immotility. Chemical Element: mendelevium. Painfulness: sharpness. Ability: midas touch. Legends: tango, bill.

Siədəɛɪtɒɛtʃrɛiədztæy

Pronounced: siuhduhayitouaytshrayiuhdztay Lawfulness: licitness. Inheritance: upbringing. Immobility: immovability. Chemical Element: darmstadtium. Painfulness: sharpness. Ability: adaptability. Legends: reduction. Prophecies: bioterrorism, peekaboo. Relations: gsəhɛpsðntvʌwlɪrɪlmy (transaminase), ʃtnwgwbvhkstugiʌwəhɪ (acetic acid), ɛzɛzəzəmɛəikumsʊknðɪ (confetti).

Ɛzɛzəzəmɛəikumsʊknðɪ

Pronounced: ayzayzuhzuhmayuhikumsooknthi Lawfulness: legality. Inheritance: birthright. Immobility: inertness. Chemical Element: noble gas. Painfulness: sharpness. Ability: midas touch. Legends: paternity suit, referral. Prophecies: thickening, snafu, contempt, last rites, thrash. Relations: ʃtnwgwbvhkstugiʌwəhɪ (compensation), aɪæssætsəfrbənrʃnkpəə (vacancy rate), gsəhɛpsðntvʌwlɪrɪlmy (trisaccharide).

Ʃtnwgwbvhkstugiʌwəhɪ

Pronounced: shtnwgwbvhkstugiuwuhhi Lawfulness: licitness. Inheritance: upbringing. Immobility: immovability. Chemical Element: bohrium. Painfulness: sharpness. Ability: physical ability. Legends: cottage industry, indignity. Prophecies: application, operations, offer, prayer meeting. Relations: ɛzɛzəzəmɛəikumsʊknðɪ (ermine), aɪæssætsəfrbənrʃnkpəə (fluosilicate), siədəɛɪtɒɛtʃrɛiədztæy (mercury).

[halocantik]

After: American Journal of the College of Nutrition, 2008, 27: 677-689

 Honey for Nutrition and Health: a Review

 Stefan Bogdanov, PhD, Tomislav Jurendic, Robert Sieber, PhD, Peter Gallmann, PhD1

Swiss Bee Research Centre, Agroscope Liebefeld-Posieux Research Station ALP, Berne, Switzerland

 Key words: honey, nutrition, composition, glycemic index

     Due to the variation of botanical origin honey differs in appearance, sensory perception and composition. The main nutritional and health relevant components are carbohydrates, mainly fructose and glucose but also about 25 different oligosaccharides. Although honey is a high carbohydrate food, its glycemic index varies within a wide range from 32 to 85, depending on the botanical source. It contains small amounts of proteins, enzymes, amino acids, minerals, trace elements, vitamins, aroma compounds and polyphenols. The review covers the composition, the nutritional contribution of its components, its physiological and nutritional effects. It shows that honey has a variety of positive nutritional and health effects, if consumed at higher doses of 50 to 80 g per intake.

             1 Adress reprint requests to: Peter Gallmann, PhD, Swiss Bee Research Centre, Agroscope Liebefeld-Posieux Research Station ALP, CH-3003 Bern, Switzerland

Abbreviations: CHO = carbohydrate, GI = glycemic index, GL = glycemic load, ORAC = oxygen radical absorbance capacity; PGE = prostaglandin E; PGF = prostaglandin F, RDI

= recommended daily intake

Key teaching points:

 ·       About 95% of the honey dry matter is composed of carbohydrates, mainly fructose and glucose. 5-10 % of the total carbohydrates are oligosaccharides, in total about 25 different di- and trisaccharides.

·       The Glycemic Index of honey varies from 32 to 85, depending on the botanical source which is lower than sucrose (60 to 110). Fructose-rich honeys such as acacia honey have a low GI.

·       Besides, honey contains small amounts of proteins, enzymes, amino acids, minerals, trace elements, vitamins, aroma compounds and polyphenols.

·       Honey has been shown to possess antimicrobial, antiviral, antiparasitory, anti- inflammatory, antioxidant, antimutagenic and antitumor effects.

·       Due to its high carbohydrate content and functional properties honey is an excellent source of energy for athletes.

·       Most of the health promoting properties of honey are only achieved by application of rather high doses of honey such as 50 to 80 g per intake.

INTRODUCTION

As the only available natural sweetener honey was an important food for Homo sapiens from his very beginnings. Indeed, the relation between bees and man started as early as Stone Age [1]. In order to reach the sweet honey, man was ready to risk his life (Figure 1). The first written reference to honey, a Sumerian tablet writing, dating back to 2100-2000 BC, mentions honey’s use as a drug and an ointment [2]. In most ancient cultures honey has been used for both nutritional and medical purposes [2-5]. According to the bible, King Solomon has said: “Eat honey my son, because it is good” (Old Testament, proverb 24:13). The belief that honey is a nutrient, a drug and an ointment has been carried into our days. For a long time in human history it was an important carbohydrate source and the only largely available sweetener until industrial sugar production began to replace it after 1800 [2]. In the long human tradition honey has been used not only as a nutrient but also as a medicine [3]. An alternative medicine branch, called apitherapy, has developed in recent years, offering treatments based on honey and the other bee products against many diseases. The knowledge on this subject is compiled in various books [e.g.

6,7] or on relevant web pages such as www.apitherapy.com, www.apitherapy.org. The major use of honey in healing today is its application in the treatment of wounds, burns and infections which is not a subject of this review since it is reviewed elsewhere [8].

At present the annual world honey production is about 1.2 million tons, which is less than 1% of the total sugar production. The consumption of honey differs strongly from country to country. The major honey exporting countries China and Argentina have small annual consumption rates of 0.1 to 0.2 kg per capita. Honey consumption is higher in developed countries, where the home production does not always cover the market demand. In the European Union, which is both a major honey importer and producer, the annual consumption per capita varies from medium (0.3-0.4 kg) in Italy, France, Great Britain, Denmark and Portugal to high (1-1.8 kg) in Germany, Austria, Switzerland, Portugal, Hungary and Greece, while in countries such as USA, Canada and Australia the average per capita consumption is 0.6 to 0.8 kg/year [see http://www.apiservices.com/].

Different surveys on nutritional and health aspects of honey have been compiled [8- 13]. However, as they are not complete and comprehensive, we undertook the task to review all the available relevant sources on this topic.

  COMPOSITION

Table 1      The overall composition of honey is shown in Table 1. The carbohydrates are the main constituents, comprising about 95% of the honey dry weight. Beyond carbohydrates, honey contains numerous compounds such as organic acids, proteins, amino acids, minerals, polyphenols, vitamins and aroma compounds.

Summarising the data shown in Table 1 it can be concluded that the contribution of honey to the recommended daily intake is small. However, its importance with respect to nutrition lies in the manifold physiological effects [16]. It should be noted that the composition of honey depends greatly on the botanical origin [17], a fact that has been seldom considered in the nutritional and physiological studies.

           Table 2

Carbohydrates

The main sugars are the monosaccharides fructose and glucose. Additionally, about 25 different oligosacharides have been detected [18,19]. The principal oligosaccharides in blossom honey are the disaccharides sucrose, maltose, trehalose and turanose, as well as some nutritionally relevant ones such as panose, 1-kestose, 6-kestose and palatinose. Compared to blossom honey honeydew honey contains higher amounts of the oligosaccharides melezitose and raffinose. In the process of digestion after honey intake the principal carbohydrates fructose and glucose are quickly transported into the blood and can be utilized for energy requirements by the human body. A daily dose of 20 g honey will cover about 3% of the required daily energy (Table 2).

 Proteins, enzymes and amino acids

Honey contains roughly 0.5% proteins, mainly enzymes and free amino acids. The contribution of that fraction to human protein intake is marginal (Table 2).

The three main honey enzymes are diastase (amylase), decomposing starch or glycogen into smaller sugar units, invertase (sucrase, α-glucosidase), decomposing

sucrose into fructose and glucose, as well as glucose oxidase, producing hydrogen peroxide and gluconic acid from glucose.

          Table 3

Vitamins, minerals and trace compounds

The amount of vitamins and minerals is small and the contribution of honey to the recommended daily intake (RDI) of the different trace substances is marginal (Table 2). It is known that different unifloral honeys contain varying amounts of minerals and trace elements [26]. From the nutritional point of view chromium, manganese and selenium are important, especially for 1 to 15 years old children. The elements sulphur, boron, cobalt, fluoride, iodide, molybdenum and silicon can be important in human nutrition too, although there are no RDI values proposed for these elements (Table 3).

Honey contains 0.3-25 mg/kg choline and 0.06 to 5 mg/kg acetylcholine [12]. Choline is essential for cardiovascular and brain function as well as for cellular membrane composition and repair, while acetylcholine acts as a neurotransmitter.

 Aroma compounds, taste-building compounds and polyphenols

There is a wide variety of honeys with different tastes and colours, depending on their botanical origin [29]. The sugars are the main taste-building compounds.

Generally, honey with a high fructose content (e.g. acacia) are sweeter compared to those with high glucose concentration (e.g. rape). The honey aroma depends also on the quantity and type of acids and amino acids present. In the past decades extensive research on aroma compounds has been carried out and more than 500 different volatile compounds were identified in different types of honey. Indeed, most aroma building compounds vary in the different types of honey depending on its botanical origin [30]. Honey flavour is an important quality for its application in food industry and also a selection criterion for the consumer’s choice.

Polyphenols are another important group of compounds with respect to the appearance and the functional properties of honey. 56 to 500 mg/kg total polyphenols were found in different honey types [31,32]. Polyphenols in honey are mainly flavonoids (e.g. quercetin, luteolin, kaempferol, apigenin, chrysin, galangin), phenolic acids and phenolic acid derivatives [33]. These are compounds known to have antioxidant properties. The main polyphenols are the flavonoids, their content can vary between 60 and 460 μg/100 g of honey and was higher in samples produced during a dry season with high temperatures [34].

 Contaminants and toxic compounds

The same as any other natural food, honey can be contaminated by the environment, e.g. by heavy metals, pesticides, antibiotics etc. [35]. Generally, the contamination levels found in Europe do not present a health hazard. The main problem in recent years was the contamination by antibiotics, used against the bee brood diseases, but at present this problem seems to be under control. In the European Union antibiotics are not allowed for that purpose, and thus honey containing antibiotics is also not permitted to be traded on the market.

A few plants used by bees are known to produce nectar containing toxic substances. Diterpenoids and pyrrazolidine alkaloids are two main toxin groups relevant in nectar. Some plants of the Ericaceae family belonging to the sub-family Rhododendron, e.g. Rhododendron ponticum contain toxic polyhydroxylated cyclic hydrocarbons or diterpenoids [36]. The substances of the other toxin group, the pyrrazolidine alkaloids, found in different honey types and the potential intoxication by these substances is reviewed [37]. Cases of honey poisoning have been reported rarely in the literature and have concerned individuals from the following regions: Caucasus, Turkey, New Zealand, Australia, Japan, Nepal, South Africa, and also some countries in North and South America. Observed symptoms of such honey poisoning are vomiting, headache, stomach ache, unconsciousness, delirium, nausea and sight weakness. In general the poisonous plants are known to the local beekeepers and honey, which can possibly contain poisonous substances, is not marketed. To minimise risks of honey born poisoning in countries where plants with poisonous nectar are growing tourists are advised to buy honey in shops and not on the road and from individual beekeepers.

          Table 4

Glycemic index and fructose

The impact of carbohydrates on human health is discussed controversially, especially the understanding of how the carbohydrates of a given food affect the blood glucose level. Today, the dietary significance of carbohydrates is often indicated in terms of the glycemic index (GI). Carbohydrates with a low GI induce a small increase of glucose in blood, while those with a high GI induce a high blood glucose level. The only comprehensive data on honey GI are the one presented in Table 4, based mainly on data of different Australian honeys [38,39]. There is a

significant negative correlation between fructose content and GI, probably due to the different fructose/glucose ratios of the honey types tested. It is known that unifloral honeys have varying fructose content and fructose/glucose ratios [17]. Some honeys, e.g. acacia and yellow box, with relatively high concentration of fructose, have a lower GI than other honey types (Table 4). There was no significant correlation between GI and the other honey sugars. The GI values of 4 honeys found in one study varied between 69 and 74 [40], while in another one the value of a honey unidentified botanical origin was found to be 35 [41]. As the GI concept claims to predict the role of carbohydrates in the development of obesity [42], low GI honeys might be a valuable alternative to high GI sweeteners. In order to take into consideration the quantity of ingested food, a new term, the glycemic load, was introduced. It is calculated as follows: the GI value is multiplied by the carbohydrate content in a given portion and divided by 100. Values lower than 10 are considered low, between 10 and 20 are intermediate and above 20 belong to the category high. For an assumed honey portion of 25 g the glycemic load of most honey types is low and some types are in the intermediate range (Table 4).

The GI concept was developed to provide a numeric classification of carbohydrate foods, assuming that such data are useful in situations where the glucose tolerance is impaired. Therefore, food with a low GI should provide benefits with respect to diabetes and to the reduction of coronary heart disease [43]. The consumption of honey types with a low GI, e.g. acacia honey might have beneficial physiological effects and could be used by diabetes patients. An intake of 50 g honey of unspecified type by healthy people and diabetes patients led to smaller increases of blood insulin and glucose than the consumption of the same amounts of glucose or of a sugar mixture resembling to honey [44,45]. It was shown that consumption of honey has a favourable effect on diabetes patients, causing a significant decrease of plasma glucose [46-48]. Honey was well tolerated by patients with diabetes of unspecified type [49] and by diabetes type-2 patients [50-52]. According to recent studies, long term consumption of food with a high GI is a significant risk factor for type-2 diabetes patients [53]. However, the GI concept for the general population is still an object of discussions [54].

Fructose is the main sugar in most honey types (Table 1). A surplus consumption of fructose in today’s American diet, mainly in the form of high-fructose corn syrup, is suspected to be one of the main causes for overweight problems [55]. By reviewing

clinical studies these authors found that fructose ingestion causes a rise of de-novo lipogenesis, which has an unfavourable effect on energy regulation and on body weight. In rat feeding experiments the hypertriglyceridemic effect observed after intake of fructose does not take place after feeding of honey [56]. Compared to rats fed with fructose, honey-fed rats had higher plasma a-tocopherol levels, higher a- tocopherol/triacylglycerol ratios, lower plasma NOx concentrations and a lower susceptibility of the heart to lipid peroxidation. These data suggest a potential nutritional benefit of substituting fructose by honey in the ingested diets.

Ingestion of both honey (2 g/kg body weight) and fructose prevented the ethanol- induced transformation of erythrocytes in mice. In humans faster recovery from ethanol intoxication after honey administration has been reported while a higher ethanol elimination rate has also been confirmed [58,59].

       Table 5

                Table 6

DIFFERENT PHYSIOLOGICAL EFFECTS

Antimicrobial, antiviral and antiparasitic activity

Honey inhibits the growth of micro-organisms and fungi. The antibacterial effect of honey, mostly against gram-positive bacteria, is well documented [60-63]. Both bacteriostatic and bactericidal effects have been reported for many strains, many of them pathogenic (Table 5). Further, it was reported that honey has also been shown to inhibit Rubella virus in vitro [64], three species of the Leishmania parasite [65] and Echinococcus [66].

The antimicrobial effect of honey is due to different substances and depends on the botanical origin of honey [60-63]. The low water activity of honey inhibits bacterial growth. Honey glucose oxidase produces the antibacterial agent hydrogen peroxide [67], but the peroxide production capacity depends also on honey catalase activity [68]. There are also other non-peroxide antibacterial substances with different chemical origin, e.g. aromatic acids [69], unknown compounds with different chemical properties [63] and phenolics and flavonoids [70,71]. The low honey pH can also be responsible for the antibacterial activity [72].

Contrary to the non-peroxide activity, the peroxide one can be destroyed by heat, light and storage [63] (Table 6). These different factors had a bigger effect on the antibacterial activity of blossom honey than on honeydew honey. Thus, for optimum antibacterial activity, honey should be stored in a cool, dark place and be consumed when fresh.

                Table 7

Antioxidant effects

The term “oxidative stress” describes the lack of equilibrium between the production of free radicals and the antioxidant protective activity in a given organism. Protection against oxidation is thought to prevent some chronic diseases [73]. The oxidative modification of the lipoproteins is considered to be an important factor for the pathogenesis of arteriosclerosis [74]. Honey has been found to contain significant antioxidant activity including glucose oxidase, catalase, ascorbic acid, flavonoids, phenolic acids, carotenoid derivatives, organic acids, Maillard reaction products, amino acids and proteins [31,75-84]. The antioxidative activity of honey polyphenols can be measured in vitro by comparing the oxygen radical absorbance capacity (ORAC) with the total phenolics concentration (Table 7). There is a significant correlation between the antioxidant activity, the phenolic content of honey and the inhibition of the in vitro lipoprotein oxidation of human serum [85]. Furthermore, in a lipid peroxidation model system buckwheat honey showed a similar antioxidant activity as 1 mM α-tocopherol [83]. The influence of honey ingestion on the antioxidative capacity of plasma was tested in two studies [86,87]. In the first one, the trial persons were given maize syrup or buckwheat honeys with a different antioxidant capacity in a dose of 1.5 g/kg body weight. In comparison to the sugar control, honey caused an increase of both the antioxidant and the reducing serum capacity. In the second study humans received a diet supplemented with a daily honey serving of 1.2 g/kg body weight. Honey increased the body antioxidant agents: blood vitamin C concentration by 47%, β-carotene by 3%, uric acid by 12%, and glutathione reductase by 7% [87]. It should be borne in mind that the antioxidant activity depends on the botanical origin of honey and varies to a great extent in honeys from different botanical sources [31,77,78,88-90].

The impact of heat and storage time on the antioxidant capacity of clover and buckwheat honey was analysed recently [91]. While processing of clover honey did not significantly influence its antioxidant capacity, storage during 6 months reduced it by about 30%. After a given storage period the antioxidant capacity of processed and raw honeys was similar. In another study both antioxidant activity and brown pigment formation increased upon heat treatment and storage [92].

Antimutagenic and antitumor activity

Mutagenic substances act directly or indirectly by promoting mutations of the genetic structure. During the roasting and frying of food heterocyclic amines are formed, e.g. Trp-p-1 (3-Amino-1,4-dimethyl-5H-pyridol [4,3-b] indole). The antimutagenic activity of honeys from seven different floral sources (acacia, buckwheat, fireweed, soybean, tupelo and Christmas berry) against Trp-p-1 was tested by the Ames assay and compared to a sugar analogue as well as to individually tested simple sugars [93]. All honeys exhibited a significant inhibition of Trp-p-1 mutagenicity. Glucose and fructose were found to have a similar antimutagenic activity as honey. Nigerose, another sugar, present in honey [18,19] has an immunoprotective activity [94]. The anti-metastatic effect of honey and its possible mode of anti-tumor action was studied by the application of honey in spontaneous mammary carcinoma in methylcholanthrene-induced fibrosarcoma of CBA mice and in anaplastic colon adenocarcinoma of Y59 rats [95]. A statistically significant anti-metastatic effect was achieved by oral application of honey. These findings indicate that honey activates the immune system and honey ingestion may be advantageous with respect to cancer and metastasis prevention. In addition, it is postulated that honey given orally before tumour cell inoculation may have a decreased effect on tumour spreading. In another study of the same group the effect of honey on tumour growth, metastasising activity and induction of apoptosis and necrosis in murine tumour models (mammary and colon carcinoma) was investigated [96]. A pronounced antimetastatic effect was observed when honey was applied before tumour-cell inoculation (per oral 2 g kg-1 for mice or 1 g kg-1 for rats, once a day for 10 consecutive days).

In another study the anti-tumour effect of honey against bladder cancer was examined in vitro and in vivo in mice [97]. According to these results honey is an effective agent for inhibiting the growth of different bladder cancer cell lines (T24, RT4, 253J and MBT-2) in vitro. It is also effective when administered intralesionally or orally in the MBT-2 bladder cancer implantation mice models.

 Anti-inflammatory effects

Anti-inflammatory effects of honey in humans were studied by Al Waili and Boni [98] after ingestion of 70 g honey. The mean plasma concentration of thromboxane B(2) was reduced by 7%, 34%, and 35%, that of PGE(2) by 14%, 10%, and 19% at 1, 2,

and 3 hours, respectively, after honey ingestion. The level of PGF(2a) was decreased by 31% at 2 hours and by 14% at 3 hours after honey ingestion. At day 15, plasma concentrations of thromboxane B(2), PGE(2) and PGF(2a) decreased by 48%, 63% and 50%, respectively. The ingestion of honey decreased inflammation in an experimental model of inflammatory bowel disease in rats [99]. Honey administration is as effective as prednisolone treatment in an inflammatory model of colitis. The postulated mechanism of action is by preventing the formation of free radicals released from the inflamed tissues. The reduction of inflammation could be due to the antibacterial effect of honey or to a direct antiinflammatory effect. The latter hypothesis was supported in animal studies, where antiinflammatory effects of honey were observed in wounds with no bacterial infection [100].

 Various physiological effects

The effect of honey on the antibody production against thymus-dependent antigen in sheep red blood cells and thymus-independent antigen (Escherichia coli) in mice was studied [101]. Oral honey intake stimulates antibody production during primary and secondary immune responses against thymus-dependent and thymus- independent antigens.

In animal experiments honey showed an immunosuppressive activity [102]. This might explain why it has been hypothesised, that ingestion of honey can relieve pollen hypersensitivity.

In a study humans received a diet supplemented with a daily honey consumption of

1.2 g/kg body weight [87]. The effects observed in blood serum were an increase of monocytes (50 %), iron (20%), copper (33%), a slight increase of lymphocyte and eosinophil percentages, zinc, magnesium, hemoglobin and packed cell volume and a reduction of: ferritin (11%), immunoglobulin E (34%), aspartate transaminase (22%), alanine transaminase (18%), lactic acid dehydrogenase (41%), creatine kinase (33%) and fasting sugar (5%).

   NUTRITION AND HEALTH EFFECTS

Oral health

There is much debate whether honey is harmful to teeth. Some reports show a cariogenic effect of honey [103-106] or a much less cariogenic effect than sucrose

[107]. Due to its antibacterial activity honey ingestion inhibits the growth of bacteria, causing caries [108,109] and might induce a carioprotective effect [110,111]. It was shown that Manuka honey, a very potent antimicrobial honey, has a positive effect against dental plaque development and gingivitis [112] and can be used instead of refined sugar in the manufacture of candy [109].

According to electron microscope studies the ingestion of honey causes no erosion of tooth enamel as observed after drinking fruit juice [113]. Ten minutes after consumption of fruit juice tooth erosion was observed, while 30 minutes after honey ingestion the erosion was only very weak. This effect can be explained only partially by the calcium, phosphorous and fluoride levels of honey and other colloidal honey components might also play a role.

Summarising the different findings, it can be concluded that honey is probably not as cariogenic as other sugars and in some cases it can be carioprotective. But to be on the safe side, it is advised to clean the teeth after consumption of honey.

 Gastroenterology

According to the Muslim holy book “The Holy Hadith”, dating back to the 8th century AD prophet Mohamed recommended honey against diarrhoea [114]. Also, the Roman physician Celsus (ca. 25 AD) used honey as a cure for diarrhoea [115]. The application of honey for prevention and treatments of gastro-intestinal disorders such as peptic ulcers, gastritis, gastroenteritis has been reported in various books and publications from Eastern Europe [6,7,116-120] and from Arab countries [121].

Honey is a potent inhibitor of the causing agent of peptic ulcers and gastritis, Helicobacter pylori [122-124]. In rats honey acted against gastric ulcers experimentally induced by indomethacin and alcohol [125-128]. Honey is not involved in prostaglandin production, but it has a stimulatory effect on the sensory nerves in the stomach that respond to capsaicin [125,129]. A second mechanism of action has been proposed, postulating that this effect is due to the antioxidant properties of honey. Honey intake in rats prevented indomethacin-induced gastric lesions in rats by reducing the ulcer index, microvascular permeability, and myeloperoxidase activity of the stomach [130]. In addition, honey was found to maintain the level of non-protein sulfhydryl compounds (e.g. glutathione) in gastric tissue subjected to factors inducing ulceration [125,129,131,132]. Ingestion of dandelion honey reduced gastric juice acidity by 56% [133]. The gastric emptying of

saccharides after ingestion of honey was slower than that after ingestion of a mixture of glucose and fructose [134].

Other important effects of honey on human digestion have been linked to oligosaccharides. These honey constituents have prebiotic effects, similar to that of fructo-oligosaccharides [135,136]. The oligosaccharide panose was the most active oligosaccharide. The oligosaccharides cause an increase of bifidobacteria and lactobacilli and exert the prebiotic effect in a synergistic mode of action [137].

According to an invitro study on five bifidobacteria strains honey has a growth promoting effect similar to that of fructose and glucose oligosaccharides [138]. Unifloral honeys of sour-wood, alfalfa and sage origin stimulated the growth of five human intestinal bifidobacteria [139]. In another study honey increased both in vivo (small and large intestines of rats) and in vitro the building of Lactobacillus acidophilus and Lactobacillus plantarum, while sucrose had no effect [140].

In clinical studies with infants and children honey shortens the duration of bacterial diarrhoea and did not prolong the duration of non-bacterial diarrhoea [141].

In certain cases, consumption of relatively large amounts of honey (50 to 100 g) can lead to a mild laxative effect in individuals with insufficient absorption of honey fructose [142,143]. Fructose alone is less readily absorbed in the intestinal tract than fructose together with glucose [144]. The mild laxative properties of honey are used for the treatment of constipation in Eastern Europe [6].

Supplementation of honey in concentrations of 2, 4, 6 and 8 g/100 g protein fed to rats, improved protein and lipid digestibility [145].

 Cardiovascular health

The effects of ingestion of 75 g of natural honey compared to the same amount of artificial honey (fructose plus glucose) or glucose on plasma glucose, plasma insulin, cholesterol, triglycerides (TG), blood lipids, C-reactive proteins and homocysteine, most of them being risk factors for cardiovascular diseases, were studied in humans [47]. Elevation of insulin and C-reactive protein was significantly higher after glucose intake than after honey consumption. Glucose reduced cholesterol and low-density lipoprotein-cholesterol (LDL-C). Artificial honey slightly decreased cholesterol and LDL-C and elevated TG. Honey reduced cholesterol, LDL-C, and TG and slightly elevated high-density lipoprotein-cholesterol (HDL-C). In patients with hypertriglyceridemia, artificial honey increased TG, while honey decreased TG. In

patients with hyperlipidemia, artificial honey increased LDL-C, while honey decreased LDL-C. In diabetic patients, honey compared with dextrose caused a significantly lower rise of plasma glucose [47].

Honey can contain nitric oxide (NO) metabolites which are known indicators for cardiovascular disease risk. Increased levels of nitric oxides in honey might have a protecting function in cardiovascular diseases. Total nitrite concentration in different biological fluids from humans, including saliva, plasma, and urine was measured after ingestion of 80 g of honey [146,147]. Salivary, plasma and urinary NO metabolite concentrations showed a tendency to increase. Different honey types contained various concentrations of NO metabolites, darker or fresh honeys containing more NO metabolites than light or stored honey. After heating, NO metabolites decreased in all honey types.

Compared to fructose-fed rats, honey-fed rats had a higher plasma a-tocopherol level, and a higher a-tocopherol/triacylglycerol ratio, as well as lower plasma nitrate levels and lower susceptibility of the heart to lipid peroxidation [56].

 Infants

The application of honey in infant nutrition used to be a common recommendation during the last centuries and there are some interesting observations. Infants on a diet with honey had better blood formation and a higher weight gain than when a diet without honey was applied [148]. Honey was better tolerated by babies than sucrose

[149] and compared to a water based placebo significantly reduced the crying phases of infants [150]. Infants had a higher weight increase when fed by honey than by sucrose, and showed less throw up than the sucrose controls [151]. When infants were fed on honey rather than on sucrose an increase of haemoglobin content, a better skin colour and no digestion problems were encountered [152,153]. Infants on honey diet had a better weight increase and were less susceptible to diseases than infants fed normally or when given blood building agents [148].

The positive effects of honey in infant diet are attributed to effects on the digestion process. One possible cause is the well established effect of oligosaccharides on B. bifidus [154], see also section Gastroenterology. When fed on a mixture of honey and milk infants showed a regularly steady weight gain and had an acidophilic micro- organism flora rich in B. bifidus [155]. Another experiment with honey and milk showed that infants were suffering less frequently from diarrhoea, and their blood

contained more haemoglobin compared to those on a diet based on sucrose sweetened milk [152]. Honey fed infants had an improved calcium uptake, and lighter and thinner faeces [156].

However, there is a health concern for infants regarding the presence of Clostridium (Cl.) botulinum in honey. Since the presence of this bacterium in natural foods is ubiquitous and honey is a non sterilized packaged food from natural origin the risk of a low contamination level cannot be excluded. Spores of this bacterium can survive in honey, but they cannot build toxin. Thus, in the stomach of infants younger than one year the bacteria spores from honey can survive and theoretically build the toxin, while children older than 12 months can ingest honey without any risk. In some cases, infant botulism has been attributed to ingestion of honey [157-160]. In Germany one case of infant botulism per year is reported [160]. As a result of the reported infant botulism cases some honey packers (e.g. the British Honey Importers and Packers Association) place a warning on the honey label that “honey should not be given to infants under 12 months of age”. Recently, a scientific committee of the EU examined the hazard of Cl. botulinum in honey [161]. It has concluded that microbiological examinations of honey are necessary for controlling the spore concentration in honey, as the incidence of Cl. botulinum is relatively low and sporadic and as such tests will not prevent infant botulism. In the EU countries the health authorities have not issued a regulation for placing a warning label on honey jars.

 Athletic performance

The physiological action of gel and powdered forms of honey as a carbohydrate source for athlete performance was studied recently under controlled conditions by Kreider and coworkers [162-165]. Honey increased significantly the heart frequency and the blood glucose level during the performance [162]. It did not promote physical or psychological signs of hypoglycaemia in fasted athletes [163], or during resistance training [164]. In another trial the effect of low and high GI carbohydrate gels and honey were tested on a 64 km cycling performance [162,165]. Both high (glucose) and low GI (honey) gels increased cycling performance and the effect of honey was slightly better than the one of glucose. According to the above studies honey is well tolerated and can be an effective carbohydrate source for athletic performance.

Different health enhancing effects

A positive effect of honey on hepatitis A patients was found after ingestion of clover and rape honey, causing a decrease of the alanine aminotranferase activity (by 9 to 13 times) and a decrease of bilirubin production by 2.1 to 2.6 times [133].

Honey has a supportive effect on patients who have undergone a cancer radiation therapy by reducing the incidence of radiation mucositis. Patients with head and neck cancer treated with radiation therapy were given honey. There was a significant reduction in the symptomatic grade 3/4 mucositis among honey-treated patients compared to the controls; i.e. 20% versus 75%. The compliance of the honey-treated group of patients was better than the controls. 55% of the patients treated with honey showed no change or a positive gain in body weight compared to the controls, the majority of which lost weight [166]. Honey was administered to chemotherapy patients with neutropenia and was found to reduce the need for colony-stimulating factors [167]. Febrile neutropenia is a serious side effect of chemotherapy.

 Allergy

Honey allergy seems relatively uncommon; allergies reported can involve reactions varying from cough to anaphylaxis [145]. In this study it was reported that patients allergic to pollen are rarely allergic to honey, although there is one reported case of combined honey pollen allergy [168]. The incidence of honey allergy, reported in a group of 173 food allergy patients was 2.3% [cited in 169]. In this study the honey allergy is explained by the presence of components of bee origin.

  CONCLUSION

Due to variation of botanical origin honey differs in appearance, sensory perception and composition. It contains mainly carbohydrates. The glycemic index of honey varies from 32 to 87, depending on botanical origin and on fructose content. The main nutrition- and health relevant components are the carbohydrates, which make it an excellent energy source especially for children and sportsmen. Besides its main components, the carbohydrates fructose and glucose, honey contains also a great number of other constituents in small and trace amounts, producing numerous nutritional and biological effects: antimicrobial, antioxidant, antiviral, antiparasitic, antiinflammatory, antimutagenic, anticancer and immunosuppressive activities.

Different nutritional studies have confirmed various effects after honey ingestion, e.g.

enhanced gastroenterological and cardiovascular health. Besides, honey showed physiological effects on blood health indicators as well as effects on hepatitis A and radiation mucositis patients. However, it should be pointed out that most of these studies were based on relatively high honey intakes of 50 to 80 g. Honey compositions, and also its different biological effects, depend to a great extent on the botanical origin of honey. This fact was often not considered in the reviewed studies.

1          Figure 1: Prehistoric man gathering honey

2          A rock painting, made around 6000 BC. La Arana shekter, Bicorp, Eastern Spain.

3

4

5

6     Table 1: Honey composition (data in g/100 g) [14,15]

7            

Blossom honey

Honeydew honey

 average

min. - max.

average

min.  - max.

Water

17.2

15-20

16.3

15-20

Monosaccharides

    fructose

38.2

30-45

31.8

28-40

glucose

31.3

24-40

26.1

19-32

Disaccharides

    sucrose

0.7

0.1-4.8

0.5

0.1-4.7

others

5.0

2-8

4.0

1-6

Trisaccharides

    melezitose

<0.1

 4.0

0.3-22.0

erlose

0.8

0.5-6

1.0

0.1-6

others

0.5

0.5-1

3.0

0.1-6

Undetermined oligosaccharides

3.1

 10.1

 Total sugars

79.7

 80.5

 Minerals

0.2

0.1-0.5

0.9

0.6-2.0

Amino acids,  proteins

0.3

0.2-0.4

0.6

0.4-0.7

Acids

0.5

0.2-0.8

1.1

0.8-1.5

pH-value

3.9

3.5-4.5

5.2

4.5-6.5

8

9

10

11

1          Table 2: Honey nutrients (values compiled after different authors [14,20-27] and

2          recommended daily intake [28])

3            

Ingredient

Amount in 100 g

Recommended Daily  Intake1

  1-4

years old

4-15

years old

After 15 years  old

Energy

kcal

    Carbohydrates

kcal

300

1000-1100

1400-2700

2400-3100

Proteins

g

0.5

13-14

17-46

44-59

Fats

g

0-

-

-

Minerals

mg

    Sodium (Na)

1.6-17

300

410-550

550

Calcium (Ca)

3-31

600

700-1200

1000-1200

Potassium (K)

40-3500

1000

1400-1900

2000

Magnesium (Mg)

0.7-13

80

120-310

300-400

Phosphorus (P)

2-15

500

600-1250

700-1250

Zinc (Zn)

0.05-2

3

5-9.5

7-10

Copper (Cu)

0.02-0.6

0.5-1

0.5-1

0.5-1

Iron (Fe)

0.03-4

8

8-15

10-15

Manganese (Mn)

0.02-2

1-1.5

1.5-5

2-5

Chromium (Cr)

0.01-0.3

0.02-0.06

0.02-0.1

0.03-1.5

Selenium (Se)

0.002-0.01

0.001-0.004

0.001-0.006

0.003-0.007

Vitamins

mg

    Phyllochinon (K)

ca. 0.025

15

20-50

60-70

Thiamin (B1)

0.00-0.01

0.6

0.8-1.4

1-1.3

Riboflavin (B2)

0.01-0.02

0.7

0.9-1.6

1.2-1.5

Pyridoxin (B6)

0.01-0.32

0.4

0.5-1.4

1.2-1.6

Niacin2

0.10-0.20

7

10-18

13-17

Panthothenic acid

0.02-0.11

4

4-6

6

Ascorbic acid  (C)

2.2-2.5

60

70-100

100

4          *-only major components considered

5          1 after the German Nutrition Society [28]

6          2 Niacin equivalents: 1 mg nicotinamide = 1 mg niacin = 60 mg tryptophan (= niacin-precursor)

7

1

2     Table 3: Other trace elements in honey [14,20-27]

3          

Element

mg/100 g

Element

mg/100 g

Aluminium (Al)

0.01-2.4

Lead (Pb)*

0.001-0.03

Arsenic (As)

0.014-0.026

Lithium (Li)

0.225-1.56

Barium (Ba)

0.01-0.08

Molybdenum (Mo)

0-0.004

Boron (B)

0.05-0.3

Nickel (Ni)

0-0.051

Bromine (Br)

0.4-1.3

Rubidium (Rb)

0.040-3.5

Cadmium (Cd)*

0-0.001

Silicon (Si)

0.05-24

Chlorine (Cl)

0.4-56

Strontium (Sr)

0.04-0.35

Cobalt (Co)

0.1-0.35

Sulfur (S)

0.7-26

Floride (F)

0.4-1.34

Vanadium (V)

0-0.013

Iodide (I)

10-100

Zirconium

0.05-0.08

4     *- elements regarded as toxic, can be partially of man-made origin

5

6

7

8     Table 4: Glycemic index (GI) and glycemic load (GL) for a serving (25 g) of honey 9       [38,39]

10            

 honey

origin

Fructose

g/100 g

GI

AC

g/serving

GL (per

serving)

Acacia (black locust)*

Romania

43

32

21

7

Yellow box

Australia

46

35±4

18

6

Stringy bark

Australia

52

44±4

21

9

Red gum

Australia

35

46±3

18

8

Iron bark

Australia

34

48±3

15

7

Yapunya

Australia

42

52±5

17

9

Pure Australia

Australia

 58±6

21

12

Commercial blend

Australia

38

62±3

18

11

Salvation June

Australia

32

64±5

15

10

Commercial blend

Australia

28

72±6

13

9

Honey of unspecified origin

Canada

 87±8

21

18

average

 55

55±5

18

10

Sucrose (mean  of 10 studies)

  68±5

  Glucose

  100

  11

12     AC = available carbohydrate

1     Table 5: List of bacteria that were found to be sensitive to honey [60,61]

2            

Pathogen

Infection caused

Bacillus anthracis

anthrax

Corynebacterium diphtheriae

diphtheria

Escherichia coli

diarrhoea, septicaemia,  urinary infections, wound  infections

Haemophilus influenzae

ear infections,  meningitus, respiratory infections, sinusitis

Klebsiella pneumoniae

pneumonia

Mycobacterium tuberculosis

tuberculosis

Proteus sp.

septicaemia, urinary infections

Pseudomonas aeruginosa

urinary infections, wound infections

Salmonella sp.

diarrhoea

Salmonella cholerae-suis

septicaemia

Salmonella typhi

typhoid

Salmonella typhimurium

wound infections

Serrata marcescens

septicaemia, wound  infections

Shigella sp.

dysentery

Staphylococcus aureus

abscesses., boils,  carbuncles, impetigo, wound  infections

Streptococcus faecalis

urinary infections

Streptococcus mutans

dental carries

Streptococcus pneumoniae

ear infections,  meningitis, pneumonia, sinusitis

Streptococcus pyogenes

ear infections, impetigo,  puerperal fever, rheumatic fever,  scarlet fever, sore throat,  wound infections

Vibrio choleriae

cholera

Actinomyces pyogenes, Klebsiella pneumoniae, Nocardia asteroids, Staphylococcus aureus, Streptococcus agal., dysgal., uber

mastitis

Epidermophyton floccosum, Microsporum canis, M.. gypseum, Trichophyton rubrum,  T. tonsurans, T. mentagrophytes var. ?

tinea

diff. Escherichia coli,  Salmonella, Shigella, Vibrio, Helicobacter pylori

peptic ulcer

1          Table 6: Effect of heat, light and storage time on the antibacterial activity of honey.

2         The antibacterial activity is expressed in % of the untreated controls [63] 3

Non-peroxide activity

Peroxide activity

Storage: 15 months rt

light

dark

light

dark

Blossom honey

76

86

19

48

Honeydew honey

78

80

63

70

Heat: 15 min 70oC

   Blossom honey

86

 8

Honeydew honey

94

 78

4

5 rt = room temperature 15-20oC 6

7

8

9          Table 7. Antioxidative activity (ORAC) and total phenol content of different unifloral

10         honeys [32]

11            

 Honey type

ORAC

μmol TE/g

total phenolics GAE  mg/kg

 Buckwheat Illinois

 16.95 ± 0.76

 796 ±3 2

Buckwheat

9.81 ± 0.34

nd

Buckwheat New York

9.75 ± 0.48

456 ± 55

Buckwheat

9.34 ± 0.57

nd

Buckwheat

9.17 ± 0.63

nd

Buckwheat

7.47 ± 0.27

nd

Soy (2000)

9.49 ± 0.29

nd

Soy (1996)

8.34 ± 0.51

269 ± 22

Hawaiian Christmas berry

8.87 ± 0.33

250 ± 56

Clover (January 2000)

6.53 ± 0.70

nd

Clover (July  2000)

6.05 ± 1.00

128 ± 11

Tupelo

6.48 ± 0.37

183 ± 9

Fireweed

3.09 ± 0.27

62 ± 6

Acacia

3.00 ± 0.16

46 ± 2

12         ORAC = Oxygen radical absorbance capacity,

13         TE = Trolox equivalent, GAE = gallic acid equivalent, nd = not determined

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4                       properties of true floral and false nonfloral honeys and induced gastric ulcers. J 5               Drug Res Egypt 17:103-106, 1987.

 6          127. Gharzouli K, Amira S, Gharzouli A, Khennouf S: Gastroprotective effects of

7                       honey and glucose-fructose-sucrose-maltose mixture against ethanol-,

8                       indomethacin-, and acidified aspirin-induced lesions in the rat. Exp Toxicol 9             Pathol 54:217-221, 2002.

 10         128. Gharzouli K, Gharzouli A, Amira S, Khennouf S: Prevention of ethanol-induced

11                       gastric lesions in rats by natural honey and glucose-fructose-sucrose-maltose

12                       mixture. Pharmacol Res 43:509, 2001.

 13         129. Al Swayeh OA, Ali ATMM: Effect of ablation of capsaicin-sensitive neurons on

14                       gastric protection by honey and sucralfate. Hepato-Gastroenterol 45:297-302, 15                 1998.

 16         130. Nasuti C, Gabbianelli R, Falcioni G, Cantalamessa F: Antioxidative and

17                       gastroprotective activities of anti-inflammatory formulations derived from

18                       chestnut honey in rats. Nutr Res 26:130-137, 2006.

 19         131. Ali ATMM: Natural honey exerts its protective effects against ethanol-induced

20                       gastric lesions in rats by preventing depletion of glandular nonprotein

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 22         132. Ali ATMM: Natural honey prevents ischaemia-reperfusion-induced gastric

23                       mucosal lesions and increased vascular permeability in rats. Eur J

24                       Gastroenterol Hepatol 9:1101-1107, 1997.

 25         133. Baltuskevicius A, Laiskonis A, Vysniauskiene D, Ceksteryte V, Racys J: Use of

26                       different kinds of honey for hepatitis A treatment and for reduction of increased

27                       acidity of gastric juice. Zemdirbyste, Mokslo Darbai 76:173-180, 2001.

 28         134. Pokorn D, Vukmirovic V: Velocity of gastric emptying of saccharides after

29                       administering honey and pure invert sugar, III International Apitherapy,

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 32         135. Sanz ML, Polemis N, Morales V, Corzo N, Drakoularakou A, Gibson GR, Rastall

33                       RA: In vitro investigation into the potential prebiotic activity of honey

34                       oligosaccharides. J Agric Food Chem 53:2914-2921, 2005.

 35         136. Yun YW: Fructooligosaccharides - occurrence, preparation and application.

36                       Enzyme Microb Technol 19:107-117, 1996.

 37         137. Ustunol Z: The effect of honey on the growth of bifidobacteria. Report for the

38                       National Honey Board 1-8, 2000. http://www.honey.com/downloads/bifido.pdf,

39                       accessed on 25 May 2007.

1          138. Kajiwara S, Gandhi H, Ustunol Z: Effect of honey on the growth of and acid

2                       production by human intestinal Bifidobacterium spp: An in vitro comparison with

3                       commercial oligosaccharides and inulin. J Food Prot 65:214-218, 2002.

 4          139. Shin H.S, Ustunol Z: Carbohydrate composition of honey from different floral

5                       sources and their influence on growth of selected intestinal bacteria: An in vitro

6                       comparison. Food Res Int 38:721-728, 2005.

 7          140. Shamala TR, Jyothi YS, Saibaba P: Stimulatory effect of honey on multiplication

8                       of lactic acid bacteria under in vitro and in vivo conditions. Lett Appl Microbiol 9               30:453-455, 2000.

 10     141. Haffejee IE, Moosa A: Honey in the treatment of infantile gastroenteritis. Br Med 11          J 290:1866-1867, 1985.

 12         142. Ladas SD, Haritos DN, Raptis SA: Honey may have a laxative effect on normal

13                       subjects because of incomplete fructose absorption. Am J Clin Nutr 62:1212- 14                 1215, 1995.

 15     143. Ladas SD, Raptis S.A: Honey, fructose absorption, and the laxative effect. 16                 Nutrition 15:591-592, 1999.

 17     144. Riby JE, Fujisawa T, Kretchmer N: Fructose absorption. Am J Clin Nutr 58:748- 18          753, 1993.

 19         145. Sirnik V, Koch V, Golob T: L'influence du miel sur la digestibilité des substances

20                       nutritives chez le rat albinos. III International Apitherapy, Symposium 11-15

21             September 1978, Portoroz, Yougoslava. Bukarest: Apimondia, pp 286-290, 22             1978.

 23         146. Al-Waili NS: Identification of nitric oxide metabolites in various honeys: effects of

24                       intravenous honey on plasma and urinary nitric oxide metabolites

25                       concentrations. J Med Food 6:359-364, 2003.

 26         147. Al-Waili NS, Boni NS: Honey increased saliva, plasma, and urine content of total

27                       nitrite concentrations in normal individuals. J Med Food 7, 377-380, 2004.

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29                       Buchdruckerei A. Umiker, Biel-Madretsch, pp.3-32, 1921.

 30         149. Müller L: Der Bienenhonig in der Säuglingsernährung bei Berücksichtigung

31                       einer neuen Fertignahrung. Med Monatsschrift 10:729-732, 1956.

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33                       from De Rerum Natura to evidence-based medicine. Eur J Pediatr 160:677-678, 34                 2001.

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 37     152. Takuma DT: Honig bei der Aufzucht von Säuglingen. Monatsschrift 38               Kinderheilkunde 103:160-161, 1955.

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2                       Kinderernährung. Der Landarzt 33:250-252, 1957.

 3          154. Rivero-Urgell M, Santamaria-Orleans A: Oligosaccharides: application in infant

4                       food (review). Early Hum Dev 65:43-52, 2001.

 5         155. Hübner B: Säuglingsernährung mit Honigmilch (Nektar-Mil). Münchner Medizin 6         Wochenschrift 100:311-313, 1958.

 7     156. Bianchi EM: Honey: Its importance in children's nutrition. Amer Bee J 117:733, 8             1977.

 9          157. Cox N, Hinkle R: Infant botulism. Am Fam Physician 65:1388-1392, 2002.

 10         158. Tanzi MG, Gabay MP: Association between honey consumption and infant

11                       botulism. Pharmacotherapy 22:1479-1483, 2002.

 12         159. McMaster P, Piper S, Schell D, Gillis J, Chong A: A taste of honey. J Paediatr 13                   Child Health 36:596-597, 2000.

 14         160. Müller-Bunke H, Höck A, Schöntube M, Noack R: Säuglingsbotulismus.

15                       Monatsschrift für Kinderheilkunde 3:242-245, 2000.

 16         161. European Commission: Honey and microbiological hazards. Report European

17                       Commission of Health & Consumer Protection Directorate-General 1-40, 2002.

18                       http://ec.europa.eu/food/fs/sc/scv/out53_en.pdf, assessed 13 June 2007.

 19         162. Kreider RB, Rasmussen CJ, Lancaster SL, Kerksick C, Greenwood M: Honey:

20                       An alternative sports gel. Strength Conditioning J. 24, 50-51, 2002.

 21         163. Leutholz B, Kreider R: Optimising nutrition of exercise and sport. In Wilson, T,

22                       Temple N (ed): “Nutritional Health”. Totowa, NJ: Humana Press, pp 207-235, 23                 2001.

 24         164. Earnest C, Kreider R, Lundberg J, Rasmussen C, Cowan P, Greenwood M,

25                       Almada A: Effects of pre-exercise carbohydrate feedings on glucose and insulin

26                       responses during and after resistance exercise. J Strength Cond Res 14:361, 27                 2000.

 28         165. Earnest CP, Lancaster SL, Rasmussen CJ, Kerksick CM, Lucia A, Greenwood

29 ��                     MC, Almada AL, Cowand PA, Kreider RB: Low versus high glycemic index

30                       meals carbohydrate gel ingestion during simulated 64 km cycling time trial

31                       performance. J Strength Cond Res 18:466-472, 2004.

 32         166. Biswal BM, Zakaria A, Ahmad NM: Topical application of honey in the

33                       management of radiation mucositis. A preliminary study. Support Care Cancer 34                 11:242-248, 2003.

 35         167. Zidan J, Shetver L, Gershuny A, Abzah A, Tamam S, Stein M, Friedman E:

36                       Prevention of chemotherapy-induced neutropenia by special honey intake. Med 37                 Oncol 23:549-552, 2006.

1     168. Bousquet J, Campos J, Michel F.B: Food intolerance to honey. Allergy 39:73- 2            75, 1984.

 3          169. Helbling A, Peter C, Berchtold E, Bogdanov S, Müller U: Allergy to honey:

4                       Relation to pollen and honey bee allergy. Allergy 47:41-49, 1992. 5

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[cornley-amateur-drama-society]

Fics Posted: 2025 Q1

January

i. Can you feel it? How much I depend on you? — For 76 Kisses to Valentine's Day

Working backstage often creates near-codependent friendships. And these friendships develop little rituals and inside jokes. This is one of the little rituals that exist backstage at the Cornley Polytechnic Drama Society.

ii. If dreams can't come true, then why not pretend? — For 76 Kisses to Valentine's Day

He’s always been attuned to her, he knows, despite his best efforts. She was a magnet and he was steel. No matter how much he tried, he was always drawn to her. But surely that doesn’t mean anything, right? She’s the newest member of the Drama Society’s acting cast, and the most nervous about being on stage. For god’s sake, she could barely improvise if something changed. Of course, he’s going to be looking at her, watching her and making sure she’s alright, er, staying to the script. That must be it. There’s no way he’s in love with Vanessa. He couldn’t be.

iii. there's something so terribly tender and sweet hiding behind those teeth — For 76 Kisses to Valentine's Day

Robert kisses Vanessa much more tenderly than she would’ve expected from him, given the, well, everything about his personality.

iv. that twinkle in your eye as fireworks light the sky — For 76 Kisses to Valentine's Day

To make up for the shit he pulled all throughout A Christmas Carol, Chris ends up hosting a New Year's Eve Party for the Cast at his flat. Max doesn't think his life could be going any better right now, and he's certain his new fiancée would agree.

February

i. If these delights thy mind may move, — part of trisaccharides

All he said was that he had some materials left over from the Youth Drama Society’s Valentine’s do (of a sort). It was the Cast that swarmed him for card-making materials. It’s not like he expected the adults to be excited to make Valentine’s cards like they were in Primary all over again. Or The Cornley Amateur Drama Society make homemade Valentine’s Day cards for each other. This, somehow, doesn’t go as poorly as one might expect.

ii. Pass me the soda, would you? — For Fictober 2024, part of Snack Time with Cornley

Chris is ever-beleaguered by the cast. The Techies enjoy the show. Or Cast and Crew Meeting to Decide the Next Play: Part Two

[letsplantextractme]

Discovering Aloe Arborescens Extract Supplier Uses & Benefits

This blog covers some details about the benefits of Aloe Arborescens and help you in a way to discover its great uses. To speak of, through the uses of the supreme immune health formula, the Aloe Arborescens Extract Supplier has been commended for its superior quality and satisfactory results.

Some of the aloe benefits are:

Promotes gastrointestinal tract health

Normalizes body functions

Promotes cardiovascular health and healthy blood

Help to regulate blood sugar levels

Stimulates the elimination of dysfunctional cells

Supporting apoptosis

Supporting the release of cytokines and interleukins to activate the immune cells

Gentle cleansing of toxicity for the whole body

Rejuvenation support for serious immune system suppression

Optimal immune function fortification

A complementary and integrative health care product

Clinical studies and highlights

Aloe Arborescens contains the full array of aloe health promoting, active glucomannan like mannose to guarantee max effectiveness in immune system support

Furnishing the body complete spectrum derived from liquid made from the leaf of the plant

You can find the recipe that renders user with a better health like the way mentioned earlier. Here, the readers can help you and you can come up with more particular benefits and uses to expand on the clinical studies and highlights.

Understanding Raffinose Pentahydrate

Raffinose Pentahydrate Supplier makes use of fermentable sugar ref during in vitro colonic fermentation. The packaging makes use of biochemical and physiol actions, 25kg in poly bottle and 10mg in glass bottle.

Raffinose belonging to the family of oligosaccharides, is extracted from different plants that includes seed of food legumes, cottonseed meal and sugar molasses. Raffinose improves immune response and beet sugar molasses thereby improving healing property. Further this is used as an additive in cosmetics, food and medicine. Being a trisaccharide, it comprises glucose, fructose, galactose and is hydrolysed to D-galactose and sucrose by D-galactose.

[juniperpublishers-agriculture]

Using Xyloglucan Oligosaccharides as Biostimulant to Enhance Tobacco Tolerance to Salt Stress- Juniper Publishers

Abstract

Xyloglucan oligosaccharides (XGOs) derived from the hydrolysis of plant cell wall xyloglucan, are a new class of naturally occurring biostimulants that exert a positive effect on plant growth and morphology and can enhance plant’s tolerance to stress. Here, we aimed to determine the influence of exogenous Tamarindus indica L. cell wall-derived XGOs on Nicotiana tabacum’s tolerance to salt stress by examining the plant’s morphology, physiological, and metabolic changes after XGO application. N. tabacum plants were grown in solid media for two months under salt stress with 100mM of sodium chloride (NaCl) ± 0.1μM XGO. Germination percentage (GP), number of leaves (NL), foliar area (FA), primary root length (PRL), and density of lateral roots (DLR) were measured. Also, 21-old-day N. tabacum plants were treated with a salt shock (100mM NaCl) ± 0.1μM XGOs. Proline, total chlorophyll, and total carbonyl contents in addition to lipid peroxidation degree and activities of four enzymes related to oxidative stress were quantified. Results showed that under saline conditions, XGOs caused a significant increase in NL and PRL, promoted lateral root formation, produced an increase in proline and total Chl contents, while reducing protein oxidation and lipid peroxidation. Although they modulated the activity of the enzymes analyzed, they were not statistically different from the salt control. XGOs may act as metabolic inducers that trigger the physiological responses for counteracting the negative effects of oxidative stress under saline conditions.

Keywords: Antioxidant system; Biostimulants; Nicotiana tabacum; Salt stress; Xyloglucan oligosaccharides

Abbreviations: CAT: Catalase; Chl: Chlorophyll; DLR: Density of Lateral Roots; FA: Foliar Area; GP: Germination Percentage; GPX: Peroxidase; GR: Glutathione Reductase; MS: Murashige and Skoog; NaCl: Sodium Chloride; NL: Number of Leaves; PCA: Principal Component Analysis; PRL: Primary Root Length; RL: Lateral Roots; ROS: Reactive Oxygen Species; SOD: Superoxide Dismutase; XGOs: Xyloglucan Oligosaccharides

    Introduction

Modern agriculture faces many challenges in order to meet the growing demand for worldwide food. The world’s population is growing at an accelerated rate. By the end of 2050 it is expected to reach 9.8 billion people and 11.2 billion in 2100 according to the “World Population Prospects: The 2017 Revision”, published by the United Nations Department of Economic and Social Affairs. However, food productivity and availability are decreasing as a result of the effects of several biotic and abiotic factors. Therefore, several actions are being taken to reduce these losses and to cope with the growing food need for the world’s population.

Soil salinity is a worldwide phenomenon that occurs under almost all climatic conditions and is a major impediment to achieving increased crop yields. Using the FAO/UNESCO soil map of the world (1970–1980), FAO estimated that 19.5% of irrigated land were salt-affected soils, and of the almost 1.5 billion ha of dryland agriculture, 32 million (2.1%) suffer from salinity problems [1]. Salt-affected soils are characterized by abundant quantities of neutral soluble salts that adversely affect plant uptake of nutrients in the soil and their growth [2]. Under salt stress, plants are also under other types of stresses, which have deleterious effects on them such as water stress, ionic toxicity, and nutritional deficiencies [2]. Altogether, these conditions confer oxidative stress and metabolic imbalance to plants [3]. Consequently, plants exposed to high saline conditions shown growth inhibition or retardation.

The morphology of plants exposed to salinity can be affected by soil salt concentrations, type of plant species, age, and plant stages (vegetative or flowering), and/or the type of salt present [4,5]. For example, there is a decrease in plant lengths, leaf (foliar) areas, leaf numbers and root systems under high concentrations of NaCl [4]. Also, many studies confirm the inhibitory effects of salinity on photosynthesis by changing chlorophyll content thus affecting Chl components and damaging the photosynthetic apparatus [5].

In addition, plants exposed to high NaCl concentrations (such as100-200mM) show rapid overproduction of reactive oxygen species, which have detrimental effects on the plants’ cells. ROS causes membrane lipid component peroxidation and oxidation of cellular components such as proteins and nucleic acids, which finally lead to programmed cell death [6,7]. ROS-initiated damage is reduced and repaired by a complex antioxidant system, which combines enzymatic and non-enzymatic components. It consists of low molecular weight antioxidant metabolites, including ascorbic acid, carotenoids, glutathione, α-tocopherol and enzymes such as catalase, peroxidase, superoxide dismutase, glutathione reductase and others. The degree of cellular damage will depend on the balance between ROS production and elimination by the antioxidant scavenging system [8].

Plants also accumulate compatible solutes in response to salt stress, which provides protection to them by participating in ROS detoxification and cellular osmotic regulation in addition to contributing to enzyme/protein stabilization and membrane integrity protection [6]. Among them, proline is one of the most important ones due to its multiple roles as part of the plant’s response to various types of stresses. It functions as an osmolyte for osmotic adjustment, buffering cellular redox potential under stress conditions, maintaining protein integrity, enhancing different enzymes activities, and free radical scavenging [9,10]. Its accumulation in leaves under salt stress has been correlated with stress tolerance in many plant species, allowing them to survive under this type of stress [6].

Many efforts have been done to overcome the problems associated with high soil salinity and salt stress in plants. However, the use of traditional physical and chemical methods for environmental restoration of salt contaminated soils demand significant investment of technological and economic resources [11]. In addition to these traditional approaches, different biostimulant classes have been used to increase crop performance under salt stress and to mitigate stress-induced limitations [12-14]. A plant biostimulant is any substance or microorganism that is applied to plants with the aim of enhancing nutrition efficiency, abiotic stress tolerance and/or crop quality traits regardless of its nutrients content. By extension, they also designate commercial products containing mixtures of such substances and/or microorganisms [15]. Plant biostimulants based on natural materials have received considerable attention by both the scientific community and commercial enterprises. According to Stratistics Market Research Consulting (MRC), the Global Biostimulants Market is accounted for $1.50 billion in 2016 and is expected to grow gradually to reach $3.79 billion by 2023 due to growing importance for organic products in agricultural industries [16]. However, understanding the mechanisms by which biostimulants act is critical to their widespread use for helping plants cope in saline-affected soils.

XGOs, derived from the breakdown of xyloglucans in plant cell walls, are emerging as a new class of naturally occurring biostimulants as a result of their positive effects on plant growth and morphology [17-19]. Plant-derived XGOs are also used as biotic pesticides and seed coating agents to maintain plant freshness in addition to capsule materials for synthetic seeds [20]. Xyloglucan is the quantitatively predominant hemicellulosic polysaccharide in the primary walls, which consists of ~20% (w/w) dicot and ~5% monocot primary cell walls [21]. Its backbone is composed of a β-(1,4)-D-glucan backbone that is quasi-regularly substituted with α-D-xylosyl residues linked to glucose through the O-6 position. In many species, the backbone has a regular pattern of three substituted glucose units followed by an unsubstituted glucose residue [22]. As a variety of complex structures can be formed, a code letter for each glucosyl residue has been defined to allow for the unambiguous naming of xyloglucan oligosaccharides. For example, XGOs can be classified as the XXXG-type of the XXGG-type, in which a capital G represents a unbranched Glcp residue and a capital F represents a Glcp residue that is substituted with a fucose- containing trisaccharide [23]. Soluble XGOs can be obtained from tamarind (Tamarindus indica L.) seeds after partial digestion with cellulase. A fraction of these XGOs have been shown to have physiologically active functions in plants and oligosaccharides, also known as oligosaccharins [19,24]. Their biological properties in plants depends on the fragmented structures and their concentrations, which need to be extremely low to get a variety of effects (10–9 - 10–8M) [17-19]. Few experimental data are available concerning the use of the XGOs as plant biostimulants for mitigating the damage imposed by salt stress conditions in plants [25]. Also, each new formulation requires a new biological evaluation to ensure that the effects are beneficial, consistent, and predictable.

For all of the above, the objective of this study was to determine the biostimulating effects of application of exogenous XGO derived from T. indica L. cell walls on N. tabacum seedlings grown under saline stress conditions with special attention to their influence on plant morphology, necessary physiological and metabolic changes to overcome stress, ROS detoxification, and antioxidant capacities.

    Materials and Methods

Oligosaccharin composition and concentration used

The XGO fraction used in this work was the same formulation previously reported and tested on plants [18,26]. Briefly, XGO was extracted and purified from tamarind (T. indica L.) seeds. The predominant composition of the XGO extracts consisted of XLLG and XXLG/XLXG with lower proportions of XXXG, XXGG, and XXG oligosaccharides as classified by Fry et al. [23]. Mass spectra obtained by matrix-assisted laser desorption ionisation-time of flight (MALDI-TOF) spectrometry [27] are shown in Supplementary Table 1&1A. Relative proportions of xyloglucan oligosaccharides obtained by MALDI and high-performance anion-exchange chromatography with pulsed amperometric detection analysis were similar (data not show). The isolated XGO fraction showed no cellulase activity, and protein could not be detected. The uniformity of the XGO mixture was confirmed by gel filtration analysis through a BioGel P2 (Bio-Rad Laboratories, Inc., Hercules, CA, USA). XGOs were used at a final concentration of 0.1μM which was selected based on previous experiments as an optimal concentration for stimulating root and leaf development in N. tabacum without causing changes in plants’ chromosome number [17,28].

Plant material and general growth conditions

Botanical seeds of N. tabacum Linn. were used as the plant model. Seeds were kindly provided by Dr. Alexis Acosta Maspons from the Institute of Biotechnology of National Autonomous University of Mexico (UNAM) (Cuernavaca, Morelos, Mexico). All seeds were harvested at the same time, kept at 4 °C in the dark, and grown under the same controlled conditions. Seeds were surface- sterilized and grown on MS [29] solid medium in a growth chamber (DAIHAN Scientific, model WISD, Korea) at 23 °C in longday conditions (16h light/8h dark) and 50% relative humidity. Sowing and the way in which each treatment was applied was according to the chosen evaluation (plant and root morphology measurements and biochemical analysis) and is explained in detail for each one in this section. Each experiment was repeated to generate three biological replicates.

Germination percentage and plant morphology measurements

Disinfected seed were sowed onto magenta boxes with MSagar- media, either alone as a negative control, supplemented with XGO at 0.1μM or 100mM NaCl to induce salt stress, or a combined (both NaCl+XGO). The concentration of the NaCl solution was determined based on experimental data (data not shown) in which it was considerable biomass decrease in the presence of 100mM NaCl was demonstrated. Each treatment consisted of nine seeds per magenta box and five boxes per each biological replicate. Germination percentage was calculated 10 days after sowing. Germination criteria were considered complete germination after the embryo emerged from the seed and a whole seedling was formed [30]. For plant morphology analysis, seedlings were grown for two months, and the number of leaves was then counted. Also, leaves were harvested by cutting three of the oldest ones to measure their foliar area, which were immediately photographed under a stereomicroscope (Olympus, Model CX31-RTSF) coupled to an Infinity Analyzer camera. FA was measured using the Infinity Analyze 3 software (Lumenera) according to the manufacturer instructions. For root length measurements and lateral root primordium frequency, sterilized seeds were plated on square Petri dishes in a vertical orientation containing MS-agar-media as control or supplemented with XGO at 0.1μM or 100mM NaCl to induce salt stress or a combination of XGO+NaCl. Each treatment consisted of six seeds per Petri square dishes and four dishes per each biological replicate. All measurements were performed on 3-week old plants that were fixed for 72h as previously described and had exhibited whole root systems [18]. The total number of lateral roots, which is the sum of the number of lateral root and number of primordia, were counted directly under a stereomicroscope (Olympus, Model CX31-RTSF). The fixed plants were then placed on a slide to allow the primary root extension and measurements by taking photographs and processing the images. A microscope (Olympus, Model SZ2-ILTS) coupled to an Infinity Analyzer camera was used, and primary roots lengths were measured using the Infinity Analyze 3 (Lumenera) software according to the manufacturer instructions. Lateral root density (DLR, represented as D in the formula) per mm of primary root length (PRL, represented as L in the formula) was calculated using the following equation: D = (RL+P)/L, where RL + P is the sum of the number of lateral root and number of primordia [31].

Induction conditions for biochemical analysis

For all of the biochemical analyses, a uniform induction experiment was designed in order to analyze XGO- (alone or in combination with salt shock) induced dynamic changes in N. tabacum seedlings. Specifically, we focused on the restitution phase (stage of resistance, continuing stress) of plants’ stress response phases [32]. Consequently, seeds were first placed in magenta boxes containing MS-agar-media to allow their homogeneous growth for 21 days. After that time, four treatments were administered:

a. One mL of sterile distilled and deionized water as a negative control.

b. A solution of 100 mM NaCl for salt shock.

c. 0.1μM XGO.

d. 0.1μM XGO+100mM NaCl were added to the top of the solid media.

Sampling leaves were taken at two and five days after induction, immediately frozen in liquid nitrogen, and stored at -80 °C until their use.

Quantification of proline and photosynthetic pigment content

After two and five days of XGO application with or without salt shock using NaCl 100mM, proline and total chlorophyll (Chl a+b) contents were measured. Proline extractions and quantifications were performed as previously described [33]. Briefly, the extract was prepared by mixing 20mg of ground leaves in 1mL of 80% ethanol, sonicated for 5min, and incubated for another 20min in the dark. The mixture was centrifuged at 20,000g for 5min, and 200μL of the supernatant were added to 400μL of reaction mix (ninhydrin 1% (w/v) in acetic acid 60 % (v/v), ethanol 20% [v/v]), and heated at 95 °C for 20min. Finally, absorbance was determined at 520nm using a FLU Ostar Omega Microplate Reader (BMG LABTECH GmbH, Germany). A calibration curve was used for proline concentration quantification and expressed as μg of proline per mg of plant fresh weight. For photosynthetic pigment content quantification, the Chl a and b and total chlorophyll (Chl a+b) contents were extracted with 80 % acetone and measured as described elsewhere [34].

Protein and lipid oxidative damage

Plant leaf tissue was ground to a fine powder with liquid nitrogen to ensure sample homogenization. Protein oxidation was measured using protein carbonyl content [35]. Briefly, 100μL of sodium phosphate buffer (PBS, pH 7.8) was added to 400mg of each sample, sonicated for 20min in the dark, centrifuged at 20,000g during 20min at 4 ºC, and the supernatant was used. Protein concentration in the supernatant was measured using a BCA Quantic Pro Sigma Kit to compare carbonyl content related to total protein content. Total carbonyl content was measured using dinitrophenylhydrazine (DNPH) reagent [35].

Lipid oxidation was analyzed with a thiobarbituric acid reactive substances (TBARS) method [36]. For the assay, 200mg of ground plant sample was submerged in 1000μL of acetone and sonicated for 5min. The mixture was then incubated for 10min in the dark and centrifuged at 4 °C and 20,000g for 5min. Then, 200μL of the supernatant was added to 300μL of a reaction mix containing 2:1 of 20 % (v/v) trichloroacetic acid (TCA) and 0.67% (w/v) of thiobarbituric acid (TBA). The reaction mix was heated at 95 °C for 15min, cooled at room temperature, and centrifuged at 20,000g at 4 °C for 20min. Lipid oxidation was measured by determining the absorbance at 532nm using a FLU Ostar Omega Microplate Reader (BMG LABTECH GmbH, Germany). The methylenedianiline (MDA) standard and standard curve for the estimation of total MDA were prepared as previously described [37]. Results were indicated as A532 per gram of plant sample.

Antioxidant enzyme activities

Plant leaves collected (0.4g) were homogenized in liquid nitrogen and 100μL of PBS (pH 7.8) containing protease inhibitor (Sigma Aldrich) concentration 5X. The mixture was then sonicated for 20 min in the dark and centrifuged at 20,000g at 4 ºC for 20min, after which time the protein content was measured using a bicinchoninic acid (BCA) Sigma® kit. Enzyme activities were determined immediately. Activities of the antioxidant enzymes, CAT (EC EC 1.11.1.6), GPX (EC 1.11.1.7), GR (EC 1.6.4.2, GR), and SOD (EC 1.15.1.1, SOD) were determined. The evaluation of enzymatic activities was performed by comparing equal amounts of total protein extracts from the samples collected.

CAT activity was measured by monitoring the enzyme-induced decomposition of an H2O2 solution at 240nm and calculated as H2O2 reduced per mg of protein per min [38]. GPX activity was assayed as previously described [39], in which the reaction mixture contained potassium phosphate buffer (100nM), guaiacol (15mM, pH 6.5), H2O2 0.05 % (v/v), and 60μL of protein extract. Guaiacol oxidation was monitored at 470nm and an enzyme unit was defined as the production of 1μm of oxidized guaiacol per mg of protein per min. SOD activity was measured by adapting the previously described chromogenic assay [40] for leaf tissue protein extract analysis. Briefly, for the reaction 225μL sodium pyrophosphate (pH 8.3, 0.025M), 18.8μL (186μM) phenazine methosulfate, 56.3μL (300μM) nitroblue tetrazolium, 93.7μL distilled water, and 5μL of protein extract were mixed. To initiate the reaction, 37.5μL (780μM) nicotinamide adenine dinucleotide was added, incubated for 1.5 minutes and 187.5μL glacial acetic acid were added to stop the reaction. The chromogen was extracted by addition of 700μL n-butanol followed by incubation for 10min, centrifugation at 20,000g for 5min, and absorbance measurement at 560nm. For GR activity measurement, the reaction was started by the addition of oxidized glutathione, and the decrease in absorbance at 340nm every min over a 3min period was read [41]. GR activity corresponded to the amount of enzyme required to oxidize 1μmol min-1 of nicotinamide adenine dinucleotide phosphate. For all enzyme activity analyses, results were expressed as U mg-1 protein.

Statistical analysis

For all variables analyzed, each experiment was performed in triplicate. The data were expressed as average ± standard deviation (SD) of the three independent replicates as a measure of dispersion. For the variable NL, FA, PRL, and DLR, the data were evaluated by an analysis of variance (ANOVA) by ranks (Kruskal Wallis test) and compared using a nonparametric multiple comparison test proposed by Conover [42] because the variables evaluated did not show a normal distribution and had heterogeneous variances. The adjustment to the premises was verified through the tests of Shapiro Wilk and Levene. PCA was performed with Pearson correlation matrices to represent a two-dimensional plane of treatment effects upon the five morphological traits [43]. The values of eigenvectors higher than the mean of the minor and the major values of the component were considered as significant.

Data obtained from biochemical analyzes were processed using a factorial ANOVA using a fixed effect model, in which the factors consisted of the treatments (XGO±NaCl) and the days after each treatment (two and five days). Previously, compliance with the normality and homogeneity premises were verified through the Shapiro-Wilk and Levene tests. All statistical analyses were performed with the InfoStat program [44].

Resultst

Effect of XGO on germination and growth of Nicotiana

The effects on GP and plant and root growth of N. tabacum seedlings, grown with MS ± 0.1μM XGO, salt stress with ± 100mM NaCl, or a combination of XGO and NaCl are shown in Table 1 and Figure 1. GP was statistically similar between the negative control and 0.1μM of XGO (close to 90%). However, GP was significantly reduced to 71% and 75% with 100 mM NaCl and 0.1μMXGO+100mM NaCl, respectively, compared to the MS negative control (P<0.05). Both salt stress control (MS+NaCl) and XGO+NaCl were statistically similar. On the other hand, 0.1μM XGO significantly caused a promotion in FA and PRL in two-month-old N. tabacum seedlings compared to negative control, but no statistical differences were observed in NL or in DLR. Moreover, when XGO was combined with 100 mM NaCl, there was a significant increase in NL, PRL, and DLR (P<0.05) related to salt control, but no statistical differences were observed in FA (Table 1 & Figure 1). Thus, addition of 100 mM NaCl caused an inhibition of NL and PRL by 36.3% and 43%, respectively, in N. tabacum seedlings compared to the MS negative control. Nevertheless, the inhibitory effects of salt stress on NL and PRL were reduced to 16.5% and 15.4%, respectively, compared to untreated control when XGO was incorporated in the media.

MS: Negative control; XGO: 0.1μM); MS+NaCl: Salt stress with 100mM NaCl; XGO+NaCl: 0.1μM XGO + 100mM NaCl; GP: Germination percentage; NL: Number of leaves; FA: Foliar area (mm2); PRL: Primary root length (mm); DLR: Density of lateral root.

The PCA explained the contribution percentage of each component to the total variation with the five quantitative characters evaluated (GP, NL, FA, PRL, and DLR) and each treatment (XGO and/or salt stress) (Table 2). The first two principal components (PC 1 and 2) justified 99.3 % of the total variation. The characters with the greatest contribution to variability consisted of NL, FA, and PRL in PC 1, and DLR and GP in the PC 2. The biplot chart of first and second component showed that PRL and DLR were significantly correlated (P<0.05) in addition to NL, FA, PG, and NL with PRL (Figure 2). There is a separation between treatments with NaCl either with or without XGO application in the first principal component, and in component 2 there was a clear separation of XGO treatments from those without XGO. The higher values in PRL, DLR, and NL correspond to treatments where XGO has applied alone, compared to those where it was combined with salt stress. In the second component, DLR and GP characters showed the highest positive and negative contribution, respectively. Also, there was a well-defined separation of XGO and XGO+NaCl from MS and MS+NaCl, with the higher average DRL seen with the XGO values.

Changes in proline and total chlorophyll content

Figure 3 shows the effect of XGO and salt stress on proline and chlorophyll contents of N. tabacum leaves measured after two and five days of induction. It can be noticed that treatments without salt (MS±XGO) exhibited no significant differences in proline content after two and five days of treatment (Figure 3a). Additionally, salinity stress in N. tabacum seedlings promoted significant proline accumulation compared to untreated control at both time points. However, the results obtained XGO+NaCl application reflects a gradual and significant increase in the proline content, which is 75.98% higher than salt stress control after five days of treatment (Figure 3a).

Total chlorophyll (Chl a+b) content was significantly higher after XGO application compared to the untreated control, which reached the highest levels among all treatments at both time points (Figure 3b). Also, as expected the Chl a+b content was significantly reduced by salinity stress in N. tabacum leaves compared to the untreated control and remained constant over time (P<0.05). On the other hand, XGO combined with NaCl produced a significantly higher Chl a+b content (29.58% upper) compared to salt stress control after five days of treatment, which reached levels similar to the negative control.

Changes in protein and lipid oxidative damage

The effects of XGO and salt stress on protein oxidation, measured as total carbonyl contents, in N. tabacum leaves at two and five days after induction is shown in Figure 4a. Plants treated with XGO exhibited no significant differences in total carbonyl content related to control plants after two and five days of treatment. On the other hand, NaCl application significantly increased its content by 67.10% compared to the untreated control after five days. In contrast, at the same time point, XGO application combined with salt stress significantly reduced protein oxidation in N. tabacum leaves by 98.45% compared to the salt stress control.

Lipid peroxidation was calculated in terms of MDA content as an indicator of lipid oxidative deterioration caused by severe oxidative stress in N. tabacum leaves at two and five days after induction (Figure 4b). XGO application caused the MDA amounts to remain constant over time. On the other hand, leaves from plants exposed to salinity stress demonstrated significantly higher MDA accumulation compared to untreated control at both time points. However, XGO application and salt stress also significantly caused a reduction in lipid peroxidation by 51.88% compared to salt stress control after five days of treatment.

Changes in activities of enzymes from the antioxidant system

We also examined four oxidative stress response enzyme markers in the context of their activities. Figure 5 shows the effect of XGO application and salt stress on the activities of the antioxidant enzymes, CAT, GPX, SOD, and GR in N. tabacum leaves after two and five days of induction. The results showed that five days after treatment with XGO alone resulted in a significantly higher CAT, GPX, and SOD activities over time compared to the negative control (P <0.05) (Figure 5a-c). GR activity was not significantly affected by the application of any treatment (Figure 5d). Salt shock with 100mM NaCl induced significantly higher CAT, GPX, and SOD activities two days after induction compared to the negative control. After five days in the presence of NaCl, GPX decreased, but CAT activity remained higher (P<0.05) than the untreated control. There was also an apparent increase in CAT, GPX and SOD activities after five days of treatment with XGO+NaCl, but their levels were statistically similar to those of salt control (Figure 5a–c).

Discussion

This study provides evidence concerning the effects of exogenous application of XGOs on enhancement of N. tabacum growth and development under saline conditions and salt shock in order to allow them to overcome the salt-stress limitations. Clearly, under the conditions used in the current work, XGO alone had a positive effect on NL, FA, and PRL although it was ineffective in promoting DRL. However, combined with continuous salt stress, this effect is rearranged, and XGO causes an increase in NL and PRL in addition to promoting lateral root formation although no changes in FA were observed compared to the salt control. Consequently, in the presence of XGO the plants managed to recover from limitations imposed by salt, so they display visible morphological characteristic improvement (Figure 1).

The analysis of eigenvalues corresponding to morphological changes is based on PCA analysis and explains >99% of the total experimental variability within the two first components. This is almost 100 % of the experimental variability that was achieved by reducing up to two principal components. The five evaluated traits showed a high contribution in one of the two first principal components; thus, each trait was very important to explain the variability observed in the experiment. PC 1, which explains the highest percent of variability (90.7%) allowed for separation of treatments under salt stress (100mM of NaCl) from those without NaCl. The highest values of NL, PRL, and FA were obtained in the medium with XGO and without salt stress, which were projected in the positive quadrant of the component. This result clearly indicates that XGO in the medium enhanced PLR and increased FA and NL in tobacco seedlings. PC 2, which explains the rest of the variability (8.6%), separates the treatments with XGO from the treatments without this biostimulant. In this component, the traits that showed the highest contribution were PG and DLR. According to these findings, the presence of XGO in the medium stimulated DLR, but the highest values of PG were obtained in the MS medium without NaCl.

These results confirmed that external application of 0.1μM XGO positively influenced plant growth and morphological features even under salt stress conditions and could be correlated with auxin-like activity. It is known that at approximately 1μM, at least four different cellotetraose-based XGOs (XXXG, XXLG, XXFG, and XLLG) mimic auxin by inducing growth [45]. In a previous study, we confirmed that auxin-like activity of the same XGO fraction mix and concentration used in this work on Arabidopsis thaliana seedlings [18]. These are important findings since the ability of a biostimulant to influence plant hormonal activity is one of their many important benefits because they can exert large influences that eventually will improve their health and growth. As plant growth regulators plays an important role as chemical messengers, they alert the plants when stressful environmental conditions exist so they can initiate or increase their stress response processes [46].

In this context, XGO may be acting as a “switches” that turn on the plants for stressful situations by altering hormonal balances. Zhang and Schmidt (1999) discuss the “switch” concept and give some examples of other types of biostimulants that reinforce our evidence. In this context, the results also suggested that exogenous XGO applications could be acting as “pre-stress conditioners” [46] and their effects are manifested by improving osmotic regulation, photosynthetic efficiency, or by causing an increase in antioxidant levels. This argument is based on the results in which it was shown that 21-day-old plants exposed to external application of 0.1μM XGO had higher proline accumulation and total Chl content in addition to higher CAT activity levels, GPX, and SOD compared to untreated plants. Of more interest was their effect when applied in combination with salt shock (XGO+NaCl) compared with those treated with NaCl alone (MS+NaCl). In this case, the highest recorded proline levels in addition to higher total Chl were observed after five days of treatments. Enhancement of this antioxidant machinery could be reflected in significant protein oxidation (total carbonyl content) and lipid peroxidation reduction. Therefore, it can be seen that the XGOs help the plant to cope with the effect of saline stress via proline accumulation.

Osmotic regulation is an important mechanism for plant cellular homeostasis under saline conditions in which proline is the most common osmolyte for osmoprotection [47]. The higher accumulations of proline recorded with XGO and NaCl at five days after treatment could be correlated with stress tolerance and may participate in the stress signal influence on adaptive responses (Figure 2) [6]. Proline also contributes to stabilization of sub-cellular structures (such as membranes and proteins), and its cytoplasmic accumulation could help reduce oxidative stress-generated plant protein and membrane damage after exposure to salinity [6]. This effect can be inferred because of lower total carbonyl content levels since protein-bound carbonyls represent a marker of global protein oxidation and lipid peroxidation products as biomarkers for oxidative stress that were observed in plants treated with XGO+NaCl (Figure 3). Therefore, XGO indirectly helps the cell cope with salt stress by maintaining cellular osmotic adjustment and protein and lipid integrity.

Another important result indicated that XGO seems to mitigate negative effects on photosynthesis in stressed plants by increasing Chl content. This parameter was used because of salinity- induced increase in chlorophyllase activity with the consequent degradation of chlorophyll (at least transiently). Consequently, we can deduce that the increase in exogenously XGO-induced total Chl content enabled tobacco plants to tolerate salt-stress in addition to promoting their development and growth. Similar results with photosynthetic pigments were obtained in our lab when the same XGO fraction mix was evaluated on A. thaliana seedling growth under saline stress [48].

Regarding the activity of antioxidative enzymes, exogenous application of 0.1μM XGO increased CAT, GPX and SOD enzyme activities compared to the negative control after five days of treatment. Due to the fact that no significant XGO+NaCl effects on enzymes’ activity that were analyzed in this work were observed, it would be advisable to analyze other antioxidant enzymes to expand the analysis in addition to determining its mode of action in the enzymatic antioxidant system.

Our study revealed that XGO seem to be working as metabolic inducers that trigger the physiological responses mentioned above. Some results support that xyloglucan fragments do not penetrate the cell, but instead, it has been suggested that the existence of specific receptors on the plasma membrane, which interact with the fragments, activate a signaling cascade inside the cell [49]. However, to date, no specific candidate has been identified as a possible receptor of these molecules [19]. Also, it has been suggested that they can promote modifications or integrate into the cell wall, which can affect not only the extracellular events in the wall but also intracellular events [50]. These authors demonstrated that incubation of pea stem segments partially bisected longitudinally with a xyloglucan oligosaccharide (9mM XXXG), accelerated the cell elongation by integration of xyloglucans as they were incorporated into the cell wall and became transglycosylated by xyloglucan endotransglycosylase (XET). According to this, XGO’s effects observed in our work may also result from a signal transduction XET-mediated or -induced cell wall modification cascade but not from the oligosaccarides’ direct actions.

Conclusion

Overall, we conclude that XGO can exert beneficial impacts on tobacco plants’ stress response either through hormone-like effects, osmotic regulation, photosynthetic efficiency improvement, and increase in antioxidant levels. They promote the proline accumulation as an organic osmolyte and increase total chlorophyll content and modify some antioxidative enzymes’ activities that eventually affect development of plant roots’ growth and development under salt stress. Further in vivo studies are needed to confirm the antioxidant effect of XGOs during salt stress in crop plants as well as to unravel their mechanism of action on oxidative responses. However, according to these results, the exogenous application of XGO as biostimulant at very low concentrations could be considered an alternative for improving the growth and productivity of crops of agronomic importance under salt stress.

To know more about Journal of Agriculture Research- https://juniperpublishers.com/artoaj/index.php

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Blood Group B Trisaccharide Conjugated to Biotin

Blood Group B Trisaccharide Conjugated to Biotin Catalog number: B2012545 Lot number: Batch Dependent Expiration Date: Batch dependent Amount: 1 mg Molecular Weight or Concentration: 885.0 g/mol Supplied as: Lyophilized Powder Applications: molecular tool for various biochemical applications Storage: -20°C Keywords: Gal-a-1,3(Fuc-a-1,2)Gal-b-1-O(CH2)3NHCO(CH2)5NH-biotin, Btri-sp biot Grade:…

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Student Exploration: Dehydration Synthesis

Vocabulary: carbohydrate, chemical formula, dehydration synthesis, disaccharide, glucose, hydrolysis, monosaccharide, polysaccharide, valence

Prior Knowledge Questions (Do these BEFORE using the Gizmo.)

If you exercise on a hot day, you need to worry about dehydration. In this context, what do you think dehydration means?

Astronauts and backpackers often bring dehydrated food. What do you think dehydrated food is?

Gizmo Warm-up

What do rice, potatoes, and sugar have in common? They are all foods rich in carbohydrates. Carbohydrates are an important energy source for your body. The basic building block of most carbohydrate compounds is the molecule glucose. Using the Dehydration Synthesis Gizmo™, you will learn about the structure of a glucose molecule and how glucose molecules can be joined together to make larger carbohydrate molecules.

To begin, select the CREATE GLUCOSE tab.

Look at the chemical formula for glucose. How many carbon (C), hydrogen (H), and oxygen (O) atoms are found in a molecule of glucose?      C:_______ H:__O:_______

Turn on Show chemical structure. Each black sphere represents a carbon, hydrogen, or oxygen atom. The lines connecting the spheres represent chemical bonds.

How many black spheres are in the diagram? _______

How does this relate to the number of carbon, hydrogen, and oxygen atoms in the chemical formula for glucose? •        

Activity A:

Build a glucose molecule

Get the Gizmo ready:

Be sure the CREATE GLUCOSE tab is still selected.

Introduction: Each element tends to form a certain number of chemical bonds. This value is the valence of the element. For example, a carbon atom has a valence of four.

Goal: Construct a molecule of glucose

Identify: The structure of a water molecule (H2O) can be written as H-O-H, with each dash representing a chemical bond. Count the number of bonds the oxygen and hydrogen atoms form in a water molecule.

What is the valence of oxygen? _______

What is the valence of hydrogen? _______

Build a model: Use the carbon, oxygen, and hydrogen atoms from the Atoms box to build a glucose molecule on the empty hexagon in the building region. Use the chemical structure in the lower right as a guide, and pay attention to the valence of each atom as you build.

Once you think you have correctly constructed the glucose molecule, click Check. If necessary, continue to modify your molecule until it is correct.

Make a diagram: Congratulations, you have completed a molecule of glucose! Click the Tools tab and click Screen shot to take a snapshot of your completed molecule. Right click the image, click Copy, and then paste the image into a blank document. Label the image “Glucose.”

Explain: How did the valence of each element help you determine the structure of the glucose molecule?

Make connections: Carbon forms the backbone of every major type of biological molecule, including carbohydrates, fats, proteins, and nucleic acids. How does carbon’s high valence relate to its ability to form these large and complex biomolecules?

Activity B:

Dehydration synthesis

Get the Gizmo ready:

Select the DEHYDRATION tab.

Question: What occurs when two glucose molecules bond?

Infer: Glucose is an example of a monosaccharide, the simplest type of carbohydrate. A disaccharide is made from bonding two monosaccharides together.

What do you think the prefixes mono- and di- mean? Mono-: __________ Di-: _________

Predict: Turn on Show description. Drag both glucose molecules into the building region. Observe the highlighted region. What do you think will happen to the atoms in this region when the glucose molecules bond?

Run Gizmo: Click Continue and watch the animation.

What happened?.

What was removed from the glucose molecules when they bonded to form maltose?

Infer: Based on what you have seen, create a balanced equation for the dehydration synthesis reaction. (Recall that the formula for glucose is C6H12O6.) You will have to determine the formula of maltose yourself.

Turn on Show current formula/equation to check your answer.

Summarize: Use what you have observed to explain what occurs during a dehydration synthesis reaction.

Apply: A trisaccharide is a carbohydrate made of three monosaccharides. What do you think would be the chemical formula of a trisaccharide made of three bonded glucose molecules?

Activity C:

Hydrolysis

Get the Gizmo ready:

Select the Hydrolysis tab.

Turn on Show description and Show current formula/equation.

Introduction: Carbohydrates made up of three or more bonded monosaccharides are known as polysaccharides. In a reaction known as hydrolysis, your body breaks down polysaccharides into individual monosaccharides that can be used by your cells for energy.

Question: What occurs when polysaccharides break up into monosaccharides?

Predict: Examine the polysaccharide in the building region and its chemical formula.

How many monosaccharides can form if this polysaccharide breaks up? __________

Recall the formula of glucose is C6H12O6. How many carbon, oxygen, and hydrogen atoms will you need for three glucose molecules?

What must be added to the polysaccharide in the Gizmo to get three glucose molecules?

Observe: Turn off Show current formula/equation. Drag a water molecule into the building region. Click Continue. What happened

Infer: Create a balanced equation for the hydrolysis reaction that just occurred.

Turn on Show current formula/equation to check your answer.

Observe: Turn off Show current formula/equation. Drag the second water molecule into the building region. Click Continue. What happened?

(Activity C continued on next page)

Activity C (continued from previous page)

Summarize: Now create a balanced equation for that shows the entire hydrolysis reaction. (In other words, the equation should show how the polysaccharide broke up into three separate glucose molecules.)

Turn on Show current formula/equation to check your answer.

Compare: How do hydrolysis reactions compare to dehydration synthesis reactions?

Apply: Amylose is a polysaccharide that consists of a long single chain of glucose molecules. Consider an amylose molecule with only four glucose molecules.

How many water molecules are released when the 4-glucose amylose forms? _____

do you think is the chemical formula for this amylose?

How many water molecules would be needed to break this amylose down into four glucose molecules?

Extend your thinking: Hydrolysis of the carbohydrates you eat begins in your mouth as you chew. How do you think this process might be affected if a person’s salivary glands were unable to produce saliva, which is mostly composed of water?

[hadescavedish]

I love those bizarre words of sugar: homopolysaccharide, heteropolysaccharide, monosaccharide, trisaccharide and tetrasaccharide.

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