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Plant
division
Plants range enormously in size Yet, all have fundamentally the same body plan Plants may be: Herbaceous Die back at end of growing season Lack tough structural (woody) parts Woody Remain standing at the end of the growing season Have structural components made of lignified cellulosic cell walls
Annuals are herbaceous Grow Reproduce Die in one year or less Biennials Take two years to complete life cycles Perennials Can be herbaceous or woody Have capacity to live more than two years Emergent growth may die back but roots, other underground growth remains alive but usually dormant (i.e. in a low state of metabolic activity)
Shoots
A Flowering Plant
The basic structure of a plant is shown Note distinguishing features, both above and below ground This is Arabidopsis thaliana, a member of the mustard family, an important model research organism
Plant tissues
Tissues are groups of cells that form a structural and functional unit Simple tissues have one cell type Complex tissues have two or more cell types Vascular plants have three tissue systems Ground tissue Energy transformation, storage and support Vascular tissue Water and nutrient conduction system Dermal tissue Covering for the plant body
Plant Organs
Plant organs are functional units that contain all three tissue systems Roots, stems, leaves, flower components and fruit are all organs They are continuous; e.g. - all vascular tissues are continuous in a plant
Cell Walls
Cell wall organization Growing plant cells produce a primary cell wall, which stretches as the cell grows A secondary cell wall may then be produced, inside the primary wall Strong, thick Secondary cell walls set limits to cell growth Cell walls are: Structural Provide defense against invading pathogens Provide signal transduction pathways for cells
Parenchyma Is an active, living tissue Simple tissue (one cell type) Thin cell walls Is evident as softer parts of a plant; the pith of a herbaceous plant; the edible part of fruit such as apples or a tuber such as a potato Some are photosynthetic Storage for important biological molecules starch, oil droplets, water Storage for plant hormones, resin, enzyme Can differentiate into other cell types such as during wound healing
Simple plant tissue Provides support in nonwoody plant organs Elongated Alive at maturity Walls are unevenly thickened, very thick in corners not uniformly distributed, usually appear as strands
Sclerenchyma Tissue
Cells have both primary and secondary cell walls Hardened walls are particularly thick Not flexible At maturity, cells are often dead Two types of cells Sclereids: hard cells found in nuts Fibers: long tapered cells that occur in patches or clumps
Cell walls contain cellulose, hemicellulose, pectin and lignin Cellulose is the major component
Most abundant polymer in the world ~40-60% of dry weight Form strong strands of parallel chains called cellulose microfibrils Cellulose strands are cemented together by hemicellulose and pectins
Hemicellulose varies in composition Some are polymers of b-1-4 glucoses with side-chains of the 5-C sugar xylose Lie flat against the cellulose fibrils Pectins are polymers of a-galacturonic acid Acid pectins are long molecules Neutral pectins act as linking molecules between acid pectins, glycoproteins and hemicellulose polymers Act as embedding glue Lignin is very hard, tough Chemistry is not fully elucidated Polymer of complex, amino-acid derived monomers Creates strong cell walls
Mainly composed of cellulose Small amounts of pectin and hemicellulose Much more pectin, esp. in thickened regions
Contain relatively large quantities of lignin, in addition to pectin, hemicellulose and cellulose
Vascular Tissues
Xylem conducts water and nutrients from roots to the rest of the plant Four different cell types Tracheids Dead at maturity Main water-conducting cells of fern and gymnosperms Relatively few plasmodesmata in end of cell in pit fields, thin areas w/only primary cell wall Vessel elements Dead at maturity stacked on top of each other Broader than tracheids, large holes in ends of cell Much more efficient at water transport Xylem parenchyma Provides storage Fibers Structural
Phloem
Transports food materials Provides structural support Complex tissue: four cell types Sieve tube cells Alive at maturity Extensive holes via sieve plate at cell ends, with shared cytoplasm between cells Reduced cellular organelle volume some lose nucleus Provide for support of movement of dissolved substances through the plant Companion cells Nucleated, support cells for sieve tubes Connected by many plasmodesmata to sieve tube cell Involved in sugar transport to sieve tube cell Fibers Phloem parenchyma
Stem Structure
Parenchyma Collenchyma
Trichome
Vascular cambium
External phloem
Dermal Cells
Two complex tissues types: epidermis and peridermis Epidermis (right) Is only type found in herbaceous plants Unspecialized live cells Usually nonphotosynthetic; clear covered with a waxy cuticle Cells tightly connected (right); prevents water loss
Stomata
Stomata are a type of pore Guard cells on either side, open usually during day; close at night and during drought
Closed Open
Guard cell
Guard cell
Trichomes
Outgrowths hairs Vary considerably in size and function Some leaf trichomes use to keep animals away Some used to eliminate salt Some thought to increase reflection to reduce stem/leaf temperature Root hairs are a type of trichome, increase surface area to increase uptake of nutrients
Meristems
Plant growth involves Cell division Cell elongation Cell differentiation Occurs locally, unlike in animals Places of growth called meristems Meristems made of cells that form new cells by mitosis Primary growth Elongation of the stem Secondary growth Increase in girth Only seen in gymnosperms and woody dicots
Elongation of the plant occurs at apical meristems, which are located at the tips of roots and shoots and within buds of stems Here, root tip is shown
Periderm
Woody plants have limitations with regard to growth The periderm is several cell layers thick, forms under the epidermis Forms outer bark of older stems and roots As the stem or root increases in diameter, the epidermis is sloughed off and replaced by periderm Complex tissue Cork cells Dead at maturity Coated with suberin, waterproof compound Cork parenchyma Storage cells
Secondary Growth
Secondary growth, the increase in girth, occurs at lateral meristems Extends along the length of the stem except at tips Vascular cambium Thin continuous cylinder within stem/root Between bark and wood Cells divide, adding more cells to the wood Cork cambium Thin cylinder of meristematic cells located in outer bark Divide to form cork cells on the outside, 1 or more layers of cork parenchyma cells on the inside Bark All tissues located outside of the vascular cambium Made of cork cells, cork cambium and cork parenchyma
Seedless Plants
From 440-410 mya (the Silurian), plants colonized land Today hundreds of thousands of species Found across all of the Earth Are complex multicellular organisms, from a few millimeters across to hundreds of feet tall. Probably arose from the charophytes, or stoneworts, a group of green algae Green algae and plants share Chlorophyll a and b, xanthophylls and carotenes Starch and cellulose Formation of cell plate during cell division Molecular data: RNA and DNA sequences from the nucleus, chloroplasts and ribosomal RNA provide evidence that charophytes are related to plants
Main issue on dry land is desiccation Plants developed a waxy cuticle to protect cellular structures Plants have stomata to allow gas exchange
In algae, gametangia (structures that release the sex cells, or gametes) are unicellular
The gametes are formed from an original single cell Only inner cells become gametes
The embryo (the new plant), develops within the female gametangium Protects the new plant from desiccation
Gametangia
Algal gametangia (a,b) are developed from a single cell that develops into the gametes Plant gametangia (c,d) are multicellular; asexual cells left behind after gamete release
Alternation of Generations
Gametophyte generation
Gives rise to haploid gametes by mitosis Gives rise to the 2n sporophyte generation when gametes fuse Undergoes meiosis to produce 1n spores Spores are first stage of the gametophyte generation
Sporophyte generation
The Gametophytes
Male structures are antheridia Produce sperm cells Female are archegonia Produce a single egg During fertilization one sperm reaches the egg, forms 2n zygote
2n zygote is first stage of the sporophyte generation Forms multicellular embryo inside the archegonium via mitosis
Sperm + egg zygote embryo
The embryo gives rise to the sporophyte, which gives rise to the sporogenous cells, which produce spores
Embryo mature sporophyte sporogenous cells
Bryophytes, seedless vascular plants, gymnosperms and flowering plants Bryophytes are seedless nonvascular plants that lack a conduction system to distribute minerals, water and nutrients
Seedless vascular plants, gymnosperms and flowering plants contain vascular tissues
Xylem for water and minerals Phloem for dissolved organics such as sugar
The relative relationships between the plants are not yet entirely clear The order of hornwort, liverwort and moss evolution not certain Here, club mosses are considered to have arisen from an extinct plant line
A Cladogram of Plants
Club mosses are thought to have arisen from a common ancestor of all vascular plant types The nonvascular plants are still considered to be more ancient than the vascular plants, all of which have lignin, a polymer that increases cell wall strength
Bryophytes
Thought to have arisen most closely from a green algal ancestor (a charophyte) Spores found in 460 million year-old sediments ~15,000 species Typically small, require damp regions because of lack of vascular system Three groups: Mosses (Phylum Bryophyta) Liverworts (picture, right) (Phylum Hepaticophyta) Hornworts (Phylum Anthocerotophyta)
~9000 species Each plant has tiny rhizoids that absorb nutrients Short stemlike structure with leaflike blades All structures are nondifferentiated simple cellular masses No roots, stems, or leaves Many mosses are dioecious; separate sexes Flagellated sperm cells swim from antheridium to archgonium via flowing water. 2n zygote grows via mitosis into multicellular sporophyte embryo; grows out of the top of the female gametophyte; turns brown at maturity Spores released from sporophyte transported by wind/water; germinate and produce a filament of cells called a protonema much like a filamentous green alga forms buds that grow into gametophytes to complete the cycle
~6000 species Flattened lobate structure (like a liver) called a thallus Not differentiated into leaves, stems or roots Leafy structures are only single layer of undifferentiated cells Both sexual and asexual reproduction Sexual: Archaegonia and antheridia on the 1n gametophyte Sometimes held on stalked structures called archegoniophores and antheridiophores which bear arcegonia and atheridia respectively Some asexual liverworts make little balls of tissue called gemmae that are born in a gemmae cup directly on the liverwort thallus Gemmae are dispersed by water droplets
Hornworts: Anthocerotophyta
~100 species Gametophytes much like liverworts Archegonia and antheridia are embedded in the gametophyte thallus After fertilization the sporophyte projects out of the gametophyte thallus; hence hornwort These may be the most ancient group of plants
Earliest vascular plant megafossils from ~420 mya ~11,000 species of ferns Also ~12 species of whisk ferns ~1000 species of club mosses ~15 species of horsetails All have vascular tissues: xylem and phloem Two types of leaves: Microphyll: Small, single vascular strand Seen only in club mosses Megaphyll: More than one vascular strand Form more typical leaves In ferns, horsetails, gymnosperms and flowering plants
Seed Plants
Seeds
Develop from
The fertilized egg cell The female gametophyte Associated tissues The embryonic sporophyte Nutritive tissue Protective coat
Consist of
The embryo is a tiny plant The spore is merely a haploid cell Allows development and growth until the new plant can establish local nutritive support Protects the developing plant against desiccation, allows extended dormant period
Seed Development
Seeds develop from an ovule held within an ovary, which is the female structure that encloses ovules
Ovule is the megasporangium Also includes the integuments, which are enclosing structures that go on to form the seed coat
Seeds may be surrounded by an ovary wall, or may be entirely naked. This serves as the major basis for phylogenetic separation of the two major groups of seed bearing plants
Gymnosperm means naked seed Gymnosperms entirely lack an ovary wall and bear naked seeds
Both have
Xylem (conduct water and mineral nutrients) Phloem (conduct dissolved sugars)
The gametophyte is not free-living It is attached to and is dependent on the sporophyte for nutrition Both gymnosperms and angiosperms are heterosporous
No. of species
Distribution
Worldwide
Worldwide
Gymnosperms
Some of the largest (sequoia and coastal redwoods) and oldest organisms (bristlecone pine) Four phyla
~235,000 species; are the dominant plants on Earth. Wide variety of sizes and forms, from small herbaceous plants to huge trees Flowers may be conspicuous or cryptic Are vascular plants that reproduce sexually by forming flowers, and after a double fertilization process , produce seeds within fruits Possess efficient water-conductive cells called vessel elements in xylem; efficient sugarconducting cells called sieve tube members in the phloem
Palms, grasses, orchids, irises, onions and lilies are all monocots Mainly herbaceous with long narrow leaves with parallel veins Flowers in threes or groups of three Monocots have a single cotyledon, or embryonic seed leaf and endosperm, a nutritive tissue in the mature seed
Oaks, roses, mustards, cacti, blueberries and sunflowers are all dicots Herbaceous or woody Typically broader leaves than monocots Flower parts usually in fours or fives Two cotyledons in the seeds
The Flower
Reproductive shoots usually on a stem; flower is referred to as the inflorescence Four parts arranged in whorls; i.e. circles sepals, petals, stamens and carpels Complete flowers have all four parts Incomplete flowers have one or more parts missing Held on a stalk called a peduncle Male parts are the stamens; female the carpels Sepals are lowermost and outermost whorl Leaf like, often green Cover flower when in bud All sepals together called the calyx Petals are the whorl above the sepals Broad, flat and thin Often brightly colored Petals referred to as the corolla
Flower Structure
The reproductive structures arise from whorls inside the petals Stamens are the structure s that hold the pollen-bearing anthers Male reproductive component Have a stalklike filament In the anthers, meiosis produces microspores that develop into pollen Each pollen grain develops into two cells one divides to produce the sperm cells, or male gametes while the other produces the pollen tube through which sperm cells travel to the ovum
The stigma, where pollen lands The style, or long structure through which pollen tube grows The ovary, which contains one or more ovules, which in turn develop into embryos when fertilized
A flower
Pistil Structure
Pistils may be simple or compound Nearly always have stigma, style and ovary structure Simple has single carpel Compound has several carpels fused together
Angiosperms are heterosporous Produce microspores and macrospores Megasporocyte in ovule undergoes meiosis to produce 4 1n megaspores 3 disintegrate one divides by mitosis to produce the female gametophyte called the embryo sac. Most commonly the embryo sac contains 7 cells with 8 1n nuclei 6 have 1 nucleus one of the six is the egg; a central one has two polar nuclei All cells but the egg and the polar nucleus cell disintegrate
Each microsporangium (pollen sac) of the anther contains many microsporocytes which undergo meiosis to form four haploid microspores Each microspore develops into an immature male gametophyte called a pollen grain Each pollen grain consists of the tube cell and the generative cell Anther splits, releases pollen Pollen transferred to female stigma and begins germination: Tube cell forms a pollen tube that grows down the style into the ovary The germinated pollen is the male gametophyte Generative cell divides to form two nonmotile sperm cells
Double fertilization
BOTH sperm participate in fertilization One sperm fuses with the egg Other fuses with the two haploid polar nuclei of the central cell to form a 3n (triploid) cell The 3n cell undergoes mitosis to generate the endosperm, which is a nutritive tissue for the seed
Double Fertilization
Each seed contains the embryo and its nutritive endosperm and is surrounded by a seed coat As seed develops, the ovary wall may thicken and entirely surround the seed(s) The ovary becomes a fruit Fruits protect the seed from desiccation and also aid in dispersal
Seed production is advantageous to longevity of the genetic material and dispersal Closed carpels that develop to make fruit aid in dispersal also via animals that eat the fruit Pollen is well-adapted to cross-fertilization via bees, bats, birds, etc Flowers attract pollinators Angiosperms have improved water and sugar transport in the xylem and phloem compared to gymnosperms and seedless vascular plants Broad leaves, well-developed roots that can store as well as collect nutrients, critical to success
The sporophyte the emergent and most obvious part of the plant, has undergone enormous adaptation, all of which aid in survival in a given region; for example: Cacti Trees Lilies Vines Shrubs
Germination
Dormancy is a state of reduced physiological activity Response to light, water, other environmental cues usually induces germination Germination is the sprouting of a seed Seed imbibes water Food stores broken down to make glucose Temperature is critical; sometimes optimal germination temperature range is very narrow Sometimes light is needed Sometimes a time delay is involved; sometimes the seed must be scarified to stimulate it to grow
Steps in Germination
Radicle (embryonic rootlet) forms, is positively geotropic (moves down into soil) Shoot appears, extends upward Dicots drag the cotyledons backwards out of the soil to protect the first foliage leaves as they extend and expand Monocots poke a coleoptile, a sharpened, wrapped bundle of sheathed cells that extend upward through soil
a. b.
Dicot: cotyledons emerge with a hook that protects first foliage leaves Monocots: cotyledon produces a pointed coleoptile
Leaves and flowers typically grow to a maximum size; this is referred to as determinate growth However, growth at apical meristems such as at root and shoot tips, are effectively endless; such growth is indeterminate
Flower when exposed to a narrow range of night length Respond to other stimuli; flowering is not affected by light length
Day-neutral
Short-Day Plants
Short-day (= long-night) Flower when night is longer than a critical period Qualitative: flower during short daylight days Quantitative: flowering accelerated by short daylight Typically flower in late summer or fall
Long-Day Plants
Long-day (= short-night) Qualitative: flower only with long daylight period Quantitative: flowering is accelerated with long daylight Flower in late spring, early summer
Animal Reproduction
Asexual Reproduction
Many animals can reproduce asexually by one parent giving rise to offspring that are genetically identical to the parent Involves splitting, budding or fragmentation of the parent Is advantageous when population density is low and mates are not readily available. Some animals reproduce both sexually and asexually Parthenogenesis is the development of an unfertilized egg into an adult Is common among some mollusks, crustaceans, insects, and some reptiles Usually parthenogenesis alternates with sexual reproduction Can be a mechanism to rapidly produce offspring when conditions are favorable Asexual reproduction is fastest way to achieve reproduction
Sexual Reproduction
Sexual reproduction is the most common type of animal reproduction Involves the fusion of two types of haploid gametes sperm and egg Male contributes a sperm; female the ovum; each provides unique genetic information Fusion of the gametes produces a zygote, which develops into a new complete organism Involves complex structure, function and behavioral adaptations
Sperm are typically small and motile; eggs are large and nonmotile External fertilization involves fusion of egg and sperm outside the body Many aquatic organisms fertilize gametes externally Release gametes simultaneously into water; requires prodigious production of gametes; meet by chance Internal fertilization involves delivery of sperm to the female body More reliable; female body provides watery medium for sperm movement and houses the developing embryo Most terrestrial animals, sharks and aquatic reptiles, birds and mammals Usually involves two different individuals Hermaphroditism is a form of sexual reproduction in which a single organism produces both eggs and sperm Some hermaphrodites are simultaneously male and female; others are first functionally one sex, then the other
If asexual reproduction is so efficient, why do most animals reproduce sexually, which is so much more complex and expensive energetically? Sexual reproduction maintains diversity in a species; by mixing genetic components and recombining them, the offspring is much more complex and able to adapt to a changing environment and may be able to better survive than either parent But what are the main issues? Possible answers: Adaptive mutations can pass through a population via sexual reproduction Perhaps sexual selection removes harmful mutations. Asexual organisms inherit all characteristics of parent Sexual organisms mix genes so can lose or at least dilute the impact of nonadaptive mutations
Animal Development?
Includes all stages in the life of an individual Here, we focus on the early stages fertilization to birth What happens during this time? Fusion of egg and sperm forms the zygote The zygote undergoes division to increase numbers of cells It becomes more complex It grows in size The cells undergo changes in gene expression and so change with time/place in the embryo; this is called differentiation Differentiation occurs via cell determination Determination is the process of commitment of a cell line to a particular developmental pathway Some cells do not change; these are stem cells
Differentiation
The ongoing differentiation results in the development of form and structure (i.e. morphogenesis) in the embryo Occurs via changes in
Cell protein expression Signaling between cells Cell migration Interactions with the extracellular matrix Controlled death (apoptosis) of cells
Fertilization
Is the union of a (usually) tiny sperm and an (usually) enormous ovum to form a zygote Determines the sex of the offspring in mammals and many other animals Stimulates changes in the egg that allow development May be divided into four steps
4.
Sperm contacts the egg Sperm or its nucleus enters the egg Egg becomes activated and developmental changes begin Sperm and egg nuclei fuse
Here, most of the discussion centers upon echinoderm fertilization; is very well understood
The plasma membrane of the sea urchin egg is surrounded by the vitelline layer and the thicker, outer jelly coat (zona pellucida in mammals) The acrosome reaction is the release of proteolytic enzymes from the acrosome of the sperm Acrosome digests a path through the external coverings If sea urchin gametes are of the same species, a protein called bindin on the acrosome adheres to a specific receptor on the vitelline membrane In sea urchins, sperm are immediately motile upon release to the sea water, due to changes in internal pH that occurs when the sperm encounter the sea water Sperm are attracted to the egg by chemotaxis In mammals, sperm must undergo capacitation, which is a maturation process which occurs in the female reproductive tract In mammals, specific interaction between sperm head proteins and zona pellucida
The microvilli of the egg membrane form a fertilization cone Sperm is drawn into the cone (right) Gamete plasma membranes fuse and the sperm is drawn into the egg cell
Aerobic respiration increases Enzyme systems become activated A burst of protein synthesis begins In most animals, the nucleus undergoes the second division of meiosis In many eggs it is possible to physically stimulate an egg to undergo activation, and even division Such parthogenetic and haploid eggs usually go through limited division series Naturally parthogenetic species have special mechanisms to retain the 2N state of their genes
Pronuclear Fusion
Once the second polar body is ejected the female pronucleus can fuse with the male pronucleus This is the genetic beginning of a new organism The haploid genetic complements of the two pronuclei form a 2N nucleus, which prepares the nucleus, and cell, for cleavage
Cleavage
During cleavage the zygote divides, giving rise to many cells The ovum contributes the majority of the zygote cytoplasm Both gametes contribute equal numbers of chromosomes Cleavage is a series of rapid mitotic division not accompanied by significant cell growth The zygote forms a two celled embryo, and continues divisions to form a ball of 32 cells called the morula The morula continues divisions to form the hollow blastula with up to several hundred cells The cells are called blastomeres The cavity of the blastula is the blastocoel
Patterns Of Cleavage
The pattern of cleavage is affected by the yolk Isolecithal eggs have a uniform yolk distribution Simple chordates and most invertebrates have isolecithal eggs Isolecithal eggs typically have holoblastic cleavage The daughter cells completely separate during cleavage Radial cleavage is typical of deuterostomes: echinoderms and Amphioxus First division is vertical; second division is at right angles Third division is horizontal at right angles to the first and forms an 8 cell embryo with 4 above and 4 cells below the last division plane Spiral cleavage is typical of protostomes: annelids and molluscs After first two divisions the plane of cleavage tilts and diagonal to the polar axis
Yolk provides energy for egg development The more metabolically active end of the cell is the animal pole, which contains less yolk Amphibian eggs contain moderate amounts of yolk; called mesolecithal Undergo holobastic cleavage but the divisions are concentrated in the animal end of the egg (below, frog)
Telolecithal Eggs
Telolecithal eggs have much yolk concentrated at the vegetal pole of the egg
Eggs of reptiles and birds are highly telolecithal Cell division takes place in the blastodisc Division is meroblastic; cells do not completely separate from each other and remain attached, at least initially, to the yolk mass In birds and some reptiles, the blastodisc splits into the epiblast (upper) and hypoblast (lower, nearest the yolk), separated by the blastocoel
Developmental Determinants
Cleavage may distribute developmental determinants in addition to changing the yolk distribution Cleavage provides building blocks for development The unequal distribution of cytoplasm of the zygote results in blastomeres with different cytoplasmic composition Mosaic development is a rigid developmental pattern Regulative development is a result of homogeneous cytoplasm, and cells produced by cleavage are equivalent Most animals have developmental patterns somewhere between these two extremes
Organogenesis
Organogenesis is the formation of the organs Arises from the layering of cells that occurs during gastrulation The layers are germ layers; they have specific fates in the developing embryo: Endoderm The innermost layer Goes on to form the gut Mesoderm In the middle Goes on to form the muscles, circulatory system, blood and many different organs Ectoderm The outermost Goes on to form the skin and nervous system
The nervous system is the first organ system to develop The notochord grows and induces overlying ectoderm to form the neural plate Cells of the neural plate fold to form the neural groove and the surrounding neural folds The neural folds fuse, forming a hollow neural tube The anterior portion forms the brain; the rest forms the spinal cord Right, neural fold formation in the human embryo
The neural crest is a critical structure that guides formation of several organ systems The neural crest forms on either side of the point of fusion Its cells migrate to form the dorsal root ganglia, the postganglionic sympathetic neurons, many sense organs and all pigment-forming cells Blocks of mesoderm called somites form on the outside of the neural tube, become the vertebrae and associated parts of the segmented body axis Other organs are stimulated to form as a result, in part of neural crest cell movements and differentiation The trachea grows from the gut and lungs develop from it The pharyngeal pouches grow laterally from the pharynx Branchial grooves meet the pharyngeal pouches and form branchial arches, important in many structures of the head The pharyngeal and branchial grooves form the gill slits and gills in aquatic vertebrates
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