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IV. Development and structures of pistil 雌蕊的发育与结构雌蕊的发育与结构 1.Composition of pistil雌蕊的组成雌蕊的组成 胚珠胚珠 融合的边缘融合的边缘 Ventral suture 柱头柱头 Carpel is modified leaf 心皮是变态的叶子心皮是变态的叶子 Pistil consists of one or more carpels and usually occupies the central position in the flower. In most angiosperms there is only one pistil in a flower comprised of one to five united carpels, but in some primitive angiosperms there may be several pistils (雌蕊)雌蕊)each comprised of a single carpel. Single pistil - from one carpel 单雌蕊单雌蕊 e.g. Fabaceae 豆科豆科 (Leguminosae) Prunus (Rosaceae) 李属(蔷薇科)李属(蔷薇科) Cercis chinensis (Redbud 紫荆紫荆)Prunus Prunus salicina (李李) Apocarpous gynoecium - from more than one simple carpel but not united 离生心皮雌蕊群离生心皮雌蕊群 e.g. Magnoliaceae 木兰科木兰科 Ranunculaceae 毛茛科毛茛科 Rosoideae 蔷薇亚科蔷薇亚科 Magnolia denudata 白玉兰白玉兰 Ranunculus japonicus 毛茛毛茛 Duchesnea indica 蛇莓蛇莓 Syncarpous pistil - from more than one carpel but united - compound pistil 合生生心皮雌蕊合生生心皮雌蕊 - 复雌蕊复雌蕊 e.g. Most Angiosperm 大多数被子植物大多数被子植物 But there are different types: Dianthus sp. 石竹属石竹属 Syncarpous pistil - from more than one carpel but united - compound pistil 合生生心皮雌蕊合生生心皮雌蕊 - 复雌蕊复雌蕊 Polygonum sp. 蓼属蓼属 Gypsophila sp. 霞草属霞草属 Alternanthera sp. 莲子草属莲子草属 2. Types of stigma柱头的类型柱头的类型 1 = stigma with multicellular trichomes(毛)(毛); 2 = ovary wall; 3 = ovary locule (子房室)(子房室); 4 = perianth with tepals (花被萼片状)(花被萼片状); 5 = ovule; 6 = embryo sac; 7 = integuments; 8 = hypostase; 9 = funiculus; 10 = receptacle Wet stigma 湿柱头湿柱头 e.g. Nicotiana 烟草属烟草属 Citrus柑橘属柑橘属 雌蕊成熟时柱头上能产生许雌蕊成熟时柱头上能产生许 多液态分泌物,多液态分泌物, Dry stigma 干柱头干柱头 Brassica油菜属油菜属 Gossypium 棉属棉属 雌蕊成熟时没有分泌物,雌蕊成熟时没有分泌物, 大多数被子植物的柱头具有乳突大多数被子植物的柱头具有乳突(papilla), 具蛋白质膜,能粘合花粉,识别作用具蛋白质膜,能粘合花粉,识别作用 A sectional view of the wet stigma of lily (Lilium sp. 百合)百合) showing a dense mat of hairs, a receptive surface for the pollen grains. The hairs secrete viscid substances (分泌黏液)分泌黏液)that are involved in the recognition of compatible pollen. If they fit together with the substances emanating from the pollen grain wall, (亲亲 合的)合的)pollen grains will germinate and produce normal pollen tubes (花粉(花粉 管)管). 1 = stigmatic hairs; 2 = ground parenchyma; 3 = stylar transmitting tissue ; 4 = stylar canal; 5 = exudate I. Development and structures of pistil 雌蕊的发育与结构雌蕊的发育与结构 3. Types of style花柱的类型花柱的类型 Solid style 实心型实心型 e.g. Gossynium 棉属棉属 Hollow style 空心型空心型 Lilium sp. 百合百合 3. Types of style花柱的类型花柱的类型 The style has solid pollen transmitting tissue (引导引导 组织)组织) O = ovary; Ovu = ovule; Sta = stamen; TrT = pollen transmitting tissue (solid); STG = stigma; STY = style Solid style 实心型实心型 e.g. Solanaceae 茄科茄科 Poaceae 禾本科禾本科 3. Types of style Hollow style 花柱的类型:花柱的类型:空心型空心型 The Lilium pistil has a hollow style. The pollen tubes grow to the ovary in contact with the glandular cells lining the stylar canal(花柱道花柱道). This pollen transmitting tissue (引导组织引导组织)consists of one layer of glandular cells, which secrete various hydrophilous and lipid substances involved in nourishment and guidance of the pollen tubes(涉及花粉管营养和引导)涉及花粉管营养和引导). 1 = stigmatic hairs; 2 = ground parenchyma; 3 = stylar transmitting tissue ; 4 = stylar canal; 5 = exudate 4. Structures of ovary 子房的结构子房的结构 1)Position of ovary 子房的位置子房的位置 There are three types of ovary positions: A) superior ovary上位子房上位子房 (e.g. tulip 郁金香郁金香) is situated on the receptacle above the points of origin of the perianth (花被)花被) parts and androecium(雄蕊群);(雄蕊群); B) inferior ovary 下位子房下位子房 (e.g. daffodil 水仙水仙 ) is situated on the top of the ovary. Calyx, corolla, and androecium fused to form a flower tube, or hypanthium (花筒)花筒); C) half-inferior ovary半下位子房半下位子房 in which the hypanthium is adnate to only the lower half of the ovary. Flower bud of cherry (Prunus virginiana 樱桃樱桃) In the cherry flower, a cup-like hypanthium is formed. However, the ovary is superior because the hypanthium does not become adnate to the ovary. 花筒花筒 花筒未与花筒未与 子房愈合,子房愈合, 仍为仍为子房子房 上位上位 花周位花周位 1)Position of ovary 子房的位置子房的位置 2) Structures of ovary 子房的结构子房的结构 Ovary wall Ovary locule Ovules Placenta(胎座)(胎座) Dorsa suture oi = outer integument) Development of ovule 胚珠的发育胚珠的发育(2) A big cell in nucellus starts development Arehesporial 孢原细胞孢原细胞 A embryo-sac mother cell, EMC 胚囊母细胞胚囊母细胞 Growth directly 水稻、小麦、百合、水稻、小麦、百合、 向日葵等向日葵等 Parietal cell 周缘细胞周缘细胞 Sporogenous cell 造孢细胞造孢细胞 crassinucellus 厚珠心厚珠心 Mitosis embryo-sac mother cell Most plants Lilium sp. 大多数被子植物是厚珠心的大多数被子植物是厚珠心的 Megasporogenesis starts with meiotic division of the diploid megasporocyte 二倍的大孢子母细胞开始减数分裂二倍的大孢子母细胞开始减数分裂 Ends with the formation of tetrad of haploid megaspores (单核胚囊)(单核胚囊). Of tetrad spores (四分体)四分体)only a chalazal one (合点那个)(合点那个)becomes functional-other three degenerate. Development of ovule 胚珠的发育胚珠的发育(3) EMC Carex sp. 莎草科苔属莎草科苔属 Process of from EMC to tetrad spores in Lilium Process of from EMC to tetrad spores in maize Types of megagametogenesis 大孢子发生的类型大孢子发生的类型 1) Polygonum type 蓼型蓼型(单孢型)单孢型) In 70% agiosperme, tetrad spores (四分体)四分体)only a chalazal one (合点端那个)(合点端那个)becomes functional Megasporo (大孢子),大孢子),other three degenerate(解体)(解体). Megasporo - 3 times of mitosis - 8 nucleus - mature (Single nuclear embryo sac embryo sac) Structure of mature embryo sac成熟胚囊的结构成熟胚囊的结构 three antipodals 反足细胞反足细胞 the central cell 中央细胞中央细胞 Synergids 助细胞助细胞 chalazal合点合点 Micropylar 珠孔珠孔 the egg Orthotropous crassinucellate bitegmic ovules in (Polygonum sp.) . 直生的双珠被、厚珠心胚珠直生的双珠被、厚珠心胚珠 O = obturator, II = inner integument, OT = outer integument, S = synergids, N = nucellus (megasporangium), E = egg cell, SN = secondary nucleus of central cell, ES = embryo sac, A = antipodals, H = hypostase, C = chalaza, VB = vascular bundle, and F = funiculus Structure of mature embryo sac - megagametophyte 成熟胚囊成熟胚囊- 雌配子体的结构雌配子体的结构 Seven cells and eight nucleus Anatropous tenuinucellate ovule in sedge 莎草的倒生薄珠心胚珠莎草的倒生薄珠心胚珠 The drawing portrays structural changes of a sedge (Carex sp.) ovule at three early developmental stages Characterization of mature Megagametophyte 成熟胚囊(雌配子体)的特化成熟胚囊(雌配子体)的特化 The antipodals (反足细胞)(反足细胞) are variable. Central cell (中央细胞)中央细胞)has a big vacuole (液泡)液泡), and polar nucleus sometime fused Into one secondary Nucleus (次生极核)次生极核). The egg cell and Synergids (助细胞)(助细胞)are highly polar. Synergids contain filiform apparatus (丝状器吸器)(丝状器吸器) 2)双孢型胚囊,)双孢型胚囊,no tetrad 3)四孢型胚囊,)四孢型胚囊,have a triploid nucleus ( antipodals and tetraploid polar nucleus) 珠孔端退化珠孔端退化 Campylotropous tenuinucellate bitegmic ovules in spiderwort (Tradescantia sp.) 紫露草的弯生薄珠心、单珠被胚珠紫露草的弯生薄珠心、单珠被胚珠 Identify: ovary wall, integuments and developing embryo sacs. Development of other embryo sacs In Calendula officinalis 金盏菊金盏菊(Asteraceae) the fully mature and ready to be doubly fertilized embryo sac is 7-celled and 8-nucleate. Jin Section Eight Anthesis, pollination and fertilization 第八节第八节开花、传粉与受精开花、传粉与受精 I Anthesis 开花开花 Concept: Smilacaceae 菝葜科植物菝葜科植物Calycanthus sp.夏腊梅夏腊梅 Blooming stage (开花期):开花期):several days to some months; Flower lasting time (一朵花持续时间)一朵花持续时间): from several min. to some months. II Pollination 传粉传粉 Concept: 1. Self-pollination 自花传粉自花传粉 e.g. wheat, rice, cotton, Citrus , peach, tomato, et al. Classical: cleistogamy 闭花受精闭花受精 e.g. pea 豌豆豌豆; barley 大麦大麦; Broad sense (广义的广义的): In crops, different flowers in same individual or even between flowers from different individuals. 2. Cross-pollination 异花传粉:异花传粉: process that pollens from one flower are transferred to stigma of another flowers. It is a evolutionary character. 是进化的特征。是进化的特征。 1) Adaptation to cross-pollination 植物对异花传粉的适应植物对异花传粉的适应 Corn (Zea) is a Monoecious (雌雄同珠)(雌雄同珠) plant 1) Adaptation to cross-pollination 植物对异花传粉的适应植物对异花传粉的适应 Stamens first mature in sunflower, pear and apple. Primula flower 报春花属报春花属 1) Adaptation to cross-pollination 植物对异花传粉的适应植物对异花传粉的适应 Primula sp. 报春花属报春花属 1) Adaptation to cross-pollination 植物对异花传粉的适应植物对异花传粉的适应 (4) Self-sterility 自花不孕自花不孕 One case is that pollens cannot germinate on the stigma, e.g. sunflower. 花粉不能萌发;花粉不能萌发; Another case is that pollen tube of self-pollen grows slowly. 花粉管生长慢花粉管生长慢 2) Medium of cross-pollination 异花传粉的媒介异花传粉的媒介 (1)Wind, - Anemophilous flowers or plants 风媒花及风媒植物风媒花及风媒植物 e.g. Gymnosperme 裸子植物;裸子植物; Poaceae 禾本科禾本科 Juglandaceae 胡桃科;胡桃科; Ulmaceae 榆科榆科 Myridaceae 杨梅科杨梅科 populus 杨属;杨属; Betula桦属桦属 Lack of perianth (花被)花被)or reduce, Filament longer, pollens small and light, and stigma (柱头)柱头)bigger. Characters of anemophilous flowers in Poaceae 禾本科风媒花特征禾本科风媒花特征 2)Medium of cross-pollination 异花传粉的媒介异花传粉的媒介 (2) Insects - entomophilous flowers or plants 虫媒花及虫媒植物虫媒花及虫媒植物 Paeonia sp. 芍药属芍药属 A bumblebee on a Aster sp. Inflorescence 紫菀属植物紫菀属植物 Characters of entomophilous flowers 虫媒花特征虫媒花特征 Beetles (甲虫)甲虫)in a flower Of Hydnora sp. Hawkmoth and Lonicera sp. 天蛾与忍冬植物天蛾与忍冬植物 Flowers are bigger and beautiful color, smell and bectary; pollen is bigger. 色、气味、蜜腺等色、气味、蜜腺等 Wasplike flower of Ophrys 像黄蜂一样的兰花像黄蜂一样的兰花 Spider and Orchids flower 蜘蛛蟹与兰花蜘蛛蟹与兰花With a large landing platform Different entomophilous flowers 各式各样的各式各样的 虫媒花虫媒花 Mint, 薄荷薄荷 Wasp and zebra orchid 黄蜂与斑马兰黄蜂与斑马兰 2)Medium of cross-pollination 异花传粉的媒介异花传粉的媒介 (3) Bird-pollinated flowers鸟媒花鸟媒花 Honeyeater and bell-fruited mallee; 蜜雀和钟果桉(澳大利亚)蜜雀和钟果桉(澳大利亚) A hummingbird and Fuchsia 蜂鸟与倒挂金钟花(柳叶菜科)蜂鸟与倒挂金钟花(柳叶菜科) 2)Medium of cross-pollination 异花传粉的媒介异花传粉的媒介 (4) mammal-pollinated flowers哺乳动物传粉植物哺乳动物传粉植物 A short-nosed bat and Bonana 短鼻蝙蝠和香蕉树短鼻蝙蝠和香蕉树 A tihy australian honey-possum And coral gum (Eucalyptus sp.) 澳大利亚蜜鼠和按树澳大利亚蜜鼠和按树 2)Medium of cross-pollination 异花传粉的媒介异花传粉的媒介 (3) water-pollinated plants水媒植物水媒植物 A aquatic ribbon weed ( Vallisneria sp.) release their flowers as “pollen boats” 水生的苦草属植物水生的苦草属植物 A large female flower Different flowers Insects eye 昆虫的眼睛昆虫的眼睛 Flowers of the evening primrose ( Oenothera, 月见草月见草) are uniform in color to the human eye,but insects see a different pattern in UV light (紫外光)紫外光). III Fertilization 受精受精 Concept: (1) Pollens germinate and the pollen tube grows Recognition and germination on stigma 在柱头上识别并萌发在柱头上识别并萌发 Generative cell divides into two sperms (精子)精子)in tube. 70% angiosperm belongs to this pattern. Three cells pollen in some plants III Fertilization 受精受精 (1) Pollens germinate and the pollen tube grows Rice, Oryza sativa III Fertilization 受精受精 (2) pollen tube grows into the ovary 花粉管进入子房花粉管进入子房 Pathway 途径途径: from micropylar mostly - porogamy 珠孔受精珠孔受精 chalazal合点合点 chalazogamy合点受精合点受精 e.g. Juglans 胡桃属胡桃属 mesogamy 中部受精中部受精 the tube into ovary between Chalazal and micropylar, e.g. Cucubita 南瓜属南瓜属 但是,最后都经助细胞进入胚囊但是,最后都经助细胞进入胚囊 Two sperms Charecters of double fertilization 双受精特点双受精特点 Triploid primary endosperm nucleus (3X初初 生胚乳核)生胚乳核) in Calendula officinalis 金盏菊金盏菊: (1)花粉管从退化助细胞丝状器进入胚囊)花粉管从退化助细胞丝状器进入胚囊 2个精细胞个精细胞助细胞先端小孔助细胞先端小孔 1个营养核个营养核释放到释放到 卵细胞与卵细胞与胼胝质塞孔胼胝质塞孔 中央细胞之间中央细胞之间阻止流动阻止流动 One sperm Another sperm 一个与卵细胞的无壁区接近并融合一个与卵细胞的无壁区接近并融合合子合子 另一个与极核接近并融合另一个与极核接近并融合zygote Finish double fertilization Significance of double fertilization 双受精意义双受精意义 双受精是被子植物所特有的现象。双受精是被子植物所特有的现象。 两个染色体单倍的精卵细胞的融合,把父、母双亲的遗传物两个染色体单倍的精卵细胞的融合,把父、母双亲的遗传物 质重新组合,形成了质重新组合,形成了兼有父母双重遗传性的合子兼有父母双重遗传性的合子; 形成了三倍体的初生胚乳核形成了三倍体的初生胚乳核,同样兼有父、母本的遗传特,同样兼有父、母本的遗传特 性,子代变异性更大,生活力更性,子代变异性更大,生活力更 强,更适应环境的变化。强,更适应环境的变化。 。 Influence of environment on pollination and fertilization 环境条件对传粉、受精的影响环境条件对传粉、受精的影响 Inner factors: 花粉败育或雄性不育、花粉粒与雌蕊柱头间花粉败育或雄性不育、花粉粒与雌蕊柱头间 的不亲和性以及植株营养不良等引起。的不亲和性以及植株营养不良等引起。 Outer factors: 气候条件、栽培措施和环境污染等。受精的气候条件、栽培措施和环境污染等。受精的 最适温度是最适温度是2630oC,最适湿度为,最适湿度为70 80% Section Nineforming, structure and development 第九节第九节 种子的形成、结构及生长发育种子的形成、结构及生长发育 After fertilization, zygote (合子)合子)develops into embryo (胚)(胚), and primary endosperm nucleus (初生胚乳核)(初生胚乳核) into Endosperm(胚乳)(胚乳), then Integument (珠被)(珠被)becomes seed coat. Two-celled embryo: Difference in Ultrastructure between terminal 2 = very thin layer of the endosperm cytoplasm surrounding the embryo; 3 = endosperm chloroplasts; 4 = upper suspensor cell 最顶端的胚柄细胞最顶端的胚柄细胞 16-celled embryo 1. development of embryo 胚的发育胚的发育 Proembryo 原胚期原胚期 Globular stage: 球形胚阶段球形胚阶段 Beginning of procambium Upper suspensor cell produces hypophysis (胚轴)胚轴) Nuclear type of endosperm development 核型胚乳发育核型胚乳发育 1. development of embryo Proembryo 原胚期原胚期 胚的发育胚的发育 1 = a portion of endosperm central vacuole; 2 = endothelium of the inner integument; 3 = embryoderm; 4 = cells of the future procambium; 5 = hypophysis; 6 = endosperm free nuclei 游离胚乳核游离胚乳核; 7 = suspensor Globular stage under EM 电镜下的球形胚阶段电镜下的球形胚阶段 Cordate (heart-shaped) embryo: cotyledons Develop 心形胚心形胚 2. Development of embryo after proembyo in Dicots 双子叶植物原胚期后,胚的发育双子叶植物原胚期后,胚的发育 Late cordate embryo, 晚心形胚晚心形胚 2. Development of embryo after proembyo in Dicots 双子叶植物原胚期后,胚的发育双子叶植物原胚期后,胚的发育 “Torpedo embryo:” 鱼雷形胚鱼雷形胚 The beginning of hypocotyl elongation 胚轴伸长胚轴伸长 Endosperm cellularization 胚乳细胞开始形成胚乳细胞开始形成 Early bending cotyledon stage of embryo development. Shoot 2 = seed coat; 3 = cotyledon; 4 = procambium; 5 = hypocotyl; 6 = pericarp (fruit wall); 7 = radicle (embryonic root); 8 = basal cell of suspensor; 9 = funiculus Diagrammatic review of Capsella (荠菜)(荠菜)embryogenesis Nicotiana (烟草)(烟草)embryogenesis: Embryo proper originates from transverse (horizontal) division of terminal cell 3. Development of embryo after proembyo in Monocots 单子叶植物原胚期后,胚的发育单子叶植物原胚期后,胚的发育 Embryogenesis in monocots: Sagittaria (泽泻)(泽泻) Single cotyledon is on top of the embryo Reduced cotyledon 3. Development of embryo after proembyo in Monocots 单子叶植物原胚期后,胚的发育单子叶植物原胚期后,胚的发育 1 3 A 2BC 34 4 5 7 8 9 6D 梨形胚梨形胚 一片子叶一片子叶 退化子叶退化子叶 4. Development of endosperm 胚乳发育胚乳发育 核型胚乳核型胚乳 (nuclear type) 细胞型胚乳细胞型胚乳(cellular type) Solanaceae 茄科茄科 沼生目胚乳沼生目胚乳(helobial type) 慈菇、独尾草属(慈菇、独尾草属(Eremurus sp.) 1 2 3 A BCD 介于核型与细胞型之间。介于核型与细胞型之间。 特点:初生胚乳核第一次分裂特点:初生胚乳核第一次分裂 后,胚囊分离成二室:珠孔室后,胚囊分离成二室:珠孔室 (较大)和合点室(较小)。(较大)和合点室(较小)。 此后,每室(主要是珠孔室)此后,每室(主要是珠孔室) 分别进行几次游离核的分裂。分别进行几次游离核的分裂。 5. prosembryum 外胚乳外胚乳 胚囊外围的珠心组织遭到破坏,最后为胚和胚乳所吸收,胚囊外围的珠心组织遭到破坏,最后为胚和胚乳所吸收, 故在多数植物成熟的种子中没有珠心组织。但少数植物的珠故在多数植物成熟的种子中没有珠心组织。但少数植物的珠 心组织始终存在,并能够随种子的发育而形成一种类似胚乳心组织始终存在,并能够随种子的发育而形成一种类似胚乳 的营养贮藏组织,称为外胚乳。的营养贮藏组织,称为外胚乳。例如,菠菜、甜菜、咖啡等成熟例如,菠菜、甜菜、咖啡等成熟 种子中就有外胚乳;种子中就有外胚乳; 6、种皮发育、种皮发育 1)双层种皮)双层种皮有些植物的胚珠具有两层珠被,其种皮也有些植物的胚珠具有两层珠被,其种皮也 可分成可分成外种皮与内种皮,外种皮与内种皮,如棉花、蓖麻、油菜等。如棉花、蓖麻、油菜等。 2)单层种皮)单层种皮有些植物的外珠被或内珠被在种子发育过有些植物的外珠被或内珠被在种子发育过 程中消失,如大豆、蚕豆、南瓜等的种皮程中消失,如大豆、蚕豆、南瓜等的种皮由外珠被发育;由外珠被发育; 而来;水稻、小麦等的种皮主要而来;水稻、小麦等的种皮主要由内珠被发育而由内珠被发育而来;玉米来;玉米 的珠被则完全解体。的珠被则完全解体。 3)有些植物的胚珠)有些植物的胚珠只有一层珠被只有一层珠被,则由该层珠被发育成,则由该层珠被发育成 种皮,如番茄、向日葵、胡桃等。种皮,如番茄、向日葵、胡桃等。 4 4)肉质种皮,如)肉质种皮,如玉兰玉兰的内珠被形成一保护层,外珠被则的内珠被形成一保护层,外珠被则 变成朱红色的肉质外种皮。变成朱红色的肉质外种皮。石榴石榴外珠被分化成坚硬的外种外珠被分化成坚硬的外种 皮,但其大部分表皮细胞呈辐射状延长成为囊状体,内含皮,但其大部分表皮细胞呈辐射状延长成为囊状体,内含 糖分和汁液,可以食用。肉质种皮在裸子植物中更为常见,糖分和汁液,可以食用。肉质种皮在裸子植物中更为常见, 如如银杏、苏铁银杏、苏铁等的种子。等的种子。 5)坚硬的种皮)坚硬的种皮 外珠被的外表皮有时会加厚并木质化,有时表皮下层也会分外珠被的外表皮有时会加厚并木质化,有时表皮下层也会分 化成骨状的石细胞。化成骨状的石细胞。 1 2 3 4 5 Section Ten Development, structure and spread 第十节第十节 果实的发育、结构及其对传播的适应果实的发育、结构及其对传播的适应 1. Structure and development of fruits 果实的发育和结构果实的发育和结构 1. Structure and development of fruits 果实的发育和结构果实的发育和结构 1. Structure and development of fruits 果实的发育和结构果实的发育和结构 1) 真果的结构真果的结构 真果的结构比较简单,外为果皮真果的结构比较简单,外为果皮(pericarp)
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