柏科植物雌球果发育的研究
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摘要
在裸子植物系统学研究中，雌球果的形态与结构是最重要的性状之一。
由于不同学者对裸子植物雌性生殖构造的形态学本质有着不同的见解，从而
导致对该类群植物系统发育研究结果的差异。柏科在裸子植物中是唯一一个
广布于南北两半球的科，前人对该科雌球果发育的研究涉及很少，为了进一
步了解柏科植物雌球果的形态学本质，并为裸子植物传粉生物学和系统学的
研究提供重要资料，本文选取了分布于北半球柏科中的7个属14个种，应
用扫描电镜、半薄切片、石蜡切片及整体透明等方法，对其雌球果各部分的
形态发生与发育过程作了详尽的研究，同时对传粉前后雌球果中的维管系统
作了深入的分析。主要结果归纳如下：
1）雌球花苞片的形态、结构和排列方式等与叶相似；但传粉后，随着苞
片基部的居间生长，其形态结构变得多样化；
2）在柏科7个属中，均未观察到种鳞发育，成熟球果的苞鳞得到充分发
育；
3）各苞片腋内的胚珠发育过程，几乎是同时发育或呈向顶发育过程；
4）在圆柏属和柏木属球果顶端的苞片腋内也有胚珠发育，其它各属雌球
花顶端的1-2对苞片腋内均没有胚珠发育；
5）在柏木属中，每一苞片腋内形成的胚珠数目较多，一般排成2-3行，
同一苞片腋内的胚珠呈向基发育，同一行胚珠为离心发育；
6）圆柏属的雌球果发育呈现多种式样，其中包括由数枚侧生胚珠过渡到
一枚顶生胚珠等情况；
7）柏科植物雌球花的维管系统在传粉前较为简单，即在苞片内仅具单一
的维管束延伸到顶端，与营养芽相似。传粉后随着苞片基部的居间生长，其
维管系统也随之复杂化。主要是在苞鳞内有新的维管束形成，一般是在苞鳞
中间维管束的两侧各发出一束维管束，这两束维管束由球果轴发出的位置略
高于中间一束，并在苞鳞基部迅速分枝，形成不同的排列式样。
成熟球果维管系统的排列方式与球果和苞鳞的形态有关，通常可分为三
类：（1）具球形球果的种类，包括圆柏属、扁柏属、柏木属和福建柏属，其
苞鳞内辐射状分枝的维管束形成两组，即一个具有水质部倒转的近轴系列和
一个维管束正常排列的远轴系列；（2）在苞鳞中只形成一组倒转的维管束，
如侧柏属和翠柏属；（3）崖柏属雌球果的苞鳞中没有倒转的维管束形成。
观察结果表明：柏科植物雌球果均为复轴性构造，其球果中胚珠着生在
球果轴上是由于苞片腋内种鳞退化的结果，因而不支持柏科植物雌球果的种
鳞与苞鳞完全合生的观点。
通过对柏科植物雌球果发育的研究结果，也为探讨松杉类植物雌球果的
演化趋势提供了重要线索。长期以来，众多研究者均认为，松杉类植物雌球
果的演化趋势是种鳞和苞鳞逐渐合并的过程，但从柏科植物雌球果苞鳞腋内
种鳞退化现象的发现表明，松杉类植物雌球果的演化趋势可能是一种种鳞逐
渐退化的过程。柏科的种鳞己完全退化，而苞鳞则相对发育。
依据雌球果发育中胚珠发育顺序及苞鳞数目等特征，认为福建柏属最原
始，而圆柏属则较为进化，其它各属则介于上述两属之间。
江泽平和王豁然（19的）提出，柏木亚科和澳洲柏亚科的主要区别在于
球果可育苞鳞的着生位置，如前者球果顶部苞鳞没有胚珠发育，而后者则是
球果顶部苞鳞腋内有胚珠发育。这一观点显然与柏木属和圆柏属的多数雌球
果顶部的苞鳞腋内有种子产生的情况不相符合。
依据本文的研究的结果及前人对柏科和相近类群的研究资料，赞同保留
独立的柏科和杉科；另外，以往侧柏曾被置于崖柏属中，但依据其雌球果苞
鳞中有倒转的维管束发育，崖柏属的崖柏和北美香柏雌球果的苞鳞中均无倒
转的维管束发育，我们支持其独立为侧柏属。
The female cone in conifer has received considerable attention not only for its systematic significance, but also for its intriguing morphological nature. In a series of detailed studies of fossil and living conifers, Florin (1954) came to the most widely accepted conclusion that the female cones of most conifers are compound structures, in which the seed scale represents a modified branch borne in the axil of a bract scale. Many researchers were in favor of that "in modern conifers, evolutionary advancement in the ovulate cone is shown by the various degrees of fusion between the ovuliferious scale and its subtending bract. Throughout the Pinaceae, the bract is only basally adnate to the scale, whereas these structures are more or less completely united in members of the Taxodiaceae and Cupressaceae."
Female cone development in some conifer species was extensively investigated, most of which supported Florin's theory. Nevertheless, the results were not consistent with the interpretation that the ovuliferious scale and bract were completely united in the female cone of Cupressaceae. The family Cupressaceae including about 20 genera, widely spread in the Northern and Southern hemispheres. As to such a large family, the relevant results derived from the limited species were not adequate for a conclusion. Therefore, it is essential to examine the female cone development in a wide range of species.
Ontogeny and vascular system of female cones from 7 northern genera of Cupressaceae (Calocedrus, Chamaecyparis, Cupressus, Fokiena, Juniperus, Platycladus, Thuja) were investigated in detail by means of SEM together with conventional sectioning technique. The main results are briefly summarized as follows:
1) The bracts were arranged in the decussated manner similar to those in leaves.
2) No, ovuliferous scale was observed in all of the species investigated.
3) In the axil of the bract, the ovules were developed simultaneously or in acropetal order.
4) In Juniperus and Cupressus, the uppermost pair of bracts were fertile or sterile. While in other genera the uppermost 1-2 pair(s) of bract were sterile.
5) In Cupressus, which have global cone with large number of ovules, 2-3 series ovules initiated in a basipetal sequence in the axil of each bract
6) Female cone development revealed the transition from several lateral ovules to the single terminal ovule in Juniperus.
7) In all the genera studied, three types of vascular system in female cones could be recognized. (1) In Juniperus, Cupressus, Chamaecyparis and Fokiena with global cone and peltate scale, radial branching of the strands formed two vascular arcs: an adaxial series of strands with inverted orientation of the xylem and an abaxial system comprising normally oriented bundles. (2) In Calocedrus and Platycladus, only a single series of inverted vascular bundles were generated in the adaxial side of the bract scale; (3) There was no inverted vascular bundle developed in the bract scale of Thuja occidentals.
The ontogeny of ovulate cone revealed that the female cone in Cupressaceae was a compound structure. In the axil of the fertile bract, the ovules were initiated on a broad swelling, which represented the aborted secondary branch system.
No ovuliferious scale was developed in Cupressaceae, which indicating that the evolutionary trends of female cone in conifers may not be the progressive fusion between the ovuliferious scale and the bract. The absence of the ovuliferious scale in Cupressaceae and some genera in Taxodiaceae implied that the ovuliferious scale had been lost from some Taxodiaceae in evolution.
Taking the evolutionary trends in the characteristics of female cone into account, Fokienia was considered as the most primitive genus, while Juniperus was regarded as the most advanced one in all the genera studied. The rest genera were considered to be intermediate between the two genera mentioned above.
Based on the characteristics of female cone in Curessaceae, together with results derived from earlier investigations, we believed
由于不同学者对裸子植物雌性生殖构造的形态学本质有着不同的见解，从而
导致对该类群植物系统发育研究结果的差异。柏科在裸子植物中是唯一一个
广布于南北两半球的科，前人对该科雌球果发育的研究涉及很少，为了进一
步了解柏科植物雌球果的形态学本质，并为裸子植物传粉生物学和系统学的
研究提供重要资料，本文选取了分布于北半球柏科中的7个属14个种，应
用扫描电镜、半薄切片、石蜡切片及整体透明等方法，对其雌球果各部分的
形态发生与发育过程作了详尽的研究，同时对传粉前后雌球果中的维管系统
作了深入的分析。主要结果归纳如下：
1）雌球花苞片的形态、结构和排列方式等与叶相似；但传粉后，随着苞
片基部的居间生长，其形态结构变得多样化；
2）在柏科7个属中，均未观察到种鳞发育，成熟球果的苞鳞得到充分发
育；
3）各苞片腋内的胚珠发育过程，几乎是同时发育或呈向顶发育过程；
4）在圆柏属和柏木属球果顶端的苞片腋内也有胚珠发育，其它各属雌球
花顶端的1-2对苞片腋内均没有胚珠发育；
5）在柏木属中，每一苞片腋内形成的胚珠数目较多，一般排成2-3行，
同一苞片腋内的胚珠呈向基发育，同一行胚珠为离心发育；
6）圆柏属的雌球果发育呈现多种式样，其中包括由数枚侧生胚珠过渡到
一枚顶生胚珠等情况；
7）柏科植物雌球花的维管系统在传粉前较为简单，即在苞片内仅具单一
的维管束延伸到顶端，与营养芽相似。传粉后随着苞片基部的居间生长，其
维管系统也随之复杂化。主要是在苞鳞内有新的维管束形成，一般是在苞鳞
中间维管束的两侧各发出一束维管束，这两束维管束由球果轴发出的位置略
高于中间一束，并在苞鳞基部迅速分枝，形成不同的排列式样。
成熟球果维管系统的排列方式与球果和苞鳞的形态有关，通常可分为三
类：（1）具球形球果的种类，包括圆柏属、扁柏属、柏木属和福建柏属，其
苞鳞内辐射状分枝的维管束形成两组，即一个具有水质部倒转的近轴系列和
一个维管束正常排列的远轴系列；（2）在苞鳞中只形成一组倒转的维管束，
如侧柏属和翠柏属；（3）崖柏属雌球果的苞鳞中没有倒转的维管束形成。
观察结果表明：柏科植物雌球果均为复轴性构造，其球果中胚珠着生在
球果轴上是由于苞片腋内种鳞退化的结果，因而不支持柏科植物雌球果的种
鳞与苞鳞完全合生的观点。
通过对柏科植物雌球果发育的研究结果，也为探讨松杉类植物雌球果的
演化趋势提供了重要线索。长期以来，众多研究者均认为，松杉类植物雌球
果的演化趋势是种鳞和苞鳞逐渐合并的过程，但从柏科植物雌球果苞鳞腋内
种鳞退化现象的发现表明，松杉类植物雌球果的演化趋势可能是一种种鳞逐
渐退化的过程。柏科的种鳞己完全退化，而苞鳞则相对发育。
依据雌球果发育中胚珠发育顺序及苞鳞数目等特征，认为福建柏属最原
始，而圆柏属则较为进化，其它各属则介于上述两属之间。
江泽平和王豁然（19的）提出，柏木亚科和澳洲柏亚科的主要区别在于
球果可育苞鳞的着生位置，如前者球果顶部苞鳞没有胚珠发育，而后者则是
球果顶部苞鳞腋内有胚珠发育。这一观点显然与柏木属和圆柏属的多数雌球
果顶部的苞鳞腋内有种子产生的情况不相符合。
依据本文的研究的结果及前人对柏科和相近类群的研究资料，赞同保留
独立的柏科和杉科；另外，以往侧柏曾被置于崖柏属中，但依据其雌球果苞
鳞中有倒转的维管束发育，崖柏属的崖柏和北美香柏雌球果的苞鳞中均无倒
转的维管束发育，我们支持其独立为侧柏属。
The female cone in conifer has received considerable attention not only for its systematic significance, but also for its intriguing morphological nature. In a series of detailed studies of fossil and living conifers, Florin (1954) came to the most widely accepted conclusion that the female cones of most conifers are compound structures, in which the seed scale represents a modified branch borne in the axil of a bract scale. Many researchers were in favor of that "in modern conifers, evolutionary advancement in the ovulate cone is shown by the various degrees of fusion between the ovuliferious scale and its subtending bract. Throughout the Pinaceae, the bract is only basally adnate to the scale, whereas these structures are more or less completely united in members of the Taxodiaceae and Cupressaceae."
Female cone development in some conifer species was extensively investigated, most of which supported Florin's theory. Nevertheless, the results were not consistent with the interpretation that the ovuliferious scale and bract were completely united in the female cone of Cupressaceae. The family Cupressaceae including about 20 genera, widely spread in the Northern and Southern hemispheres. As to such a large family, the relevant results derived from the limited species were not adequate for a conclusion. Therefore, it is essential to examine the female cone development in a wide range of species.
Ontogeny and vascular system of female cones from 7 northern genera of Cupressaceae (Calocedrus, Chamaecyparis, Cupressus, Fokiena, Juniperus, Platycladus, Thuja) were investigated in detail by means of SEM together with conventional sectioning technique. The main results are briefly summarized as follows:
1) The bracts were arranged in the decussated manner similar to those in leaves.
2) No, ovuliferous scale was observed in all of the species investigated.
3) In the axil of the bract, the ovules were developed simultaneously or in acropetal order.
4) In Juniperus and Cupressus, the uppermost pair of bracts were fertile or sterile. While in other genera the uppermost 1-2 pair(s) of bract were sterile.
5) In Cupressus, which have global cone with large number of ovules, 2-3 series ovules initiated in a basipetal sequence in the axil of each bract
6) Female cone development revealed the transition from several lateral ovules to the single terminal ovule in Juniperus.
7) In all the genera studied, three types of vascular system in female cones could be recognized. (1) In Juniperus, Cupressus, Chamaecyparis and Fokiena with global cone and peltate scale, radial branching of the strands formed two vascular arcs: an adaxial series of strands with inverted orientation of the xylem and an abaxial system comprising normally oriented bundles. (2) In Calocedrus and Platycladus, only a single series of inverted vascular bundles were generated in the adaxial side of the bract scale; (3) There was no inverted vascular bundle developed in the bract scale of Thuja occidentals.
The ontogeny of ovulate cone revealed that the female cone in Cupressaceae was a compound structure. In the axil of the fertile bract, the ovules were initiated on a broad swelling, which represented the aborted secondary branch system.
No ovuliferious scale was developed in Cupressaceae, which indicating that the evolutionary trends of female cone in conifers may not be the progressive fusion between the ovuliferious scale and the bract. The absence of the ovuliferious scale in Cupressaceae and some genera in Taxodiaceae implied that the ovuliferious scale had been lost from some Taxodiaceae in evolution.
Taking the evolutionary trends in the characteristics of female cone into account, Fokienia was considered as the most primitive genus, while Juniperus was regarded as the most advanced one in all the genera studied. The rest genera were considered to be intermediate between the two genera mentioned above.
Based on the characteristics of female cone in Curessaceae, together with results derived from earlier investigations, we believed
引文
1． Aase HC, Vascular anatomy of the megasporophylls of Conifers. Bot Gaz, 1915, 60: 277-313
2． Aulenback KR and LePage BA, Taxodium wallissi sp. nov.: first occurrence of Taxodium from the upper cretaceous. Int J Plant Sci, 1998, 159(2) : 367-390
3． Axsmith BJ, Taylor TN and Taylor EL, A new fossil conifer from the Triassic of North America: implications for models of ovulate cone scale evolution. Int J Plant Sci, 1998, 159(2) : 358-366
4． Baird AM, The life history of Callitris. Phytomorphology, 1953, 3:258-284
5． Beck CB, Origin and Evolution of Gmnosperms. New York: Columbia University Press, 1988
6． Bessey CE, The morphology of the pine cone. Bot Gaz, 1902, 33:157-159
7． Biswas C and Johri BM, Gmnosperns. Berlin: Springer-Verlag. 1997．
8． Bouman F, The ovule. In: Johri BM ed. Embryology of Angiosperms. Berlin: Springer-Verlag, 1984, 123-157
9． Brunsfeld ST Soltis PS, Soltis DE, Gadek PA, Quinn CJ, Strenge DD and Banker TM, Phylogenetic relationships among the genera of Taxodiaceae and Cupressaceae: evidence from rbcl sequences. Sys Bot, 1994, 19(2) : 253-262
10． Buchholz JT, The embryogeny of conifers. In Duggar BM ed. Proc Fourth Int Congr Plant Sci (Ithaca, New York, 1926) , Menasha, Wisconsin: Geoge Banta Publishing Company, 1929, 1:359-392
11． Buchhholz JT, The classification of Coniferales. Transaction of the Illinois State Academy of Science, 1934,25: 112-113
12． Chamberlain CJ, Gymnosperm Structure and Evolution. Chicago: The University of Chicago Press. 1935
13． Chaw SM, Zharkikh A, Sung HM et al, Molecular phylogeny of extant gymnosperm and seed plant evolution: analysis of nuclear 18S rRNA sequences. Mol Biol Evol, 1997, 14(1) : 56-68
14． Coen E and Meyerowitz EM, The war of the whorls: genetic interactions controlling flower development. Nature, 1991, 353: 31-37
15． Colombo L, Franken J, Koetje E et.al, The Petunia MADS-box gene FBP11 determines ovule identity. Plant Cell, 1995, 7: 1859-1868
16． Coulter PR and Chamberlain CJ, Morphology of Gymnosperms. Chicago: The University of Chicago Press. 1917．
17． Dalimore W and Jackson AB, A Handbook of Coniferae and Ginkgoaceae, 4th ed.
18. De Laubenfels DJ, Commentary: the organization of female fertile structure in conifers. Int J Plant Sci, 1992, 153 (4): ⅶ-ⅷ
19. De Laubenfels DJ, The relationships of Fitzroya cupressoides (Molina) Johnston and Diselma archeri J. D. Hooker based on morphological considerations. Phytomorphology, 1965, 15 (4): 414-419
20. Del Fueyo GM, Cone and ovule development in the Podocarpus species from Argentina. Phytomorphology. 1999, 49 (1): 49-60
21. Dogra PD, Embryogeny of gymnosperm and taxonomy ---- an assessment. In Nair PKK ed., Glimpses in Plant Research ---- Modern Trends in Plant Taxonomy, 1980, 5: 114-128
22. Eckenwalder JE, Comments on" A new classification of the conifers". Taxon, 1976a, 25: 337-338
23. Eckenwalder JE, Re-evaluation of Cupressaceae and Taxodiaceae: a proposed merger. Madroo, 1976b, 23 (5): 237-300
24. Endress PK, Diversity and evolutionary biology of tropical flowers. Cambridge University Press. 1994.
25. Farjon A, World Checklist and Bibliography of Conifers. The Royal Botanic Gardens, Kew. 1998
26. Florin R, Evolution in cordaites and confers. Acta Horti Bergiani, 1951, 15: 285-388
27. Florin R, The female reproductive organs of conifers and taxads. Bio Rev, 1954, 29: 367-389
28. Foster AS, Gifford EM, Comparative Morphology of Vascular Plants. New York: WH Freeman and Company, 1974.
29. Fu DZ (傅德志), Nageiaceae ---- a new gymnosperm family. Acta Phytotax Sin (植物分类学报), 1992, 30(6): 515-528
30. Fu DZ (傅德志)and Yang QE (杨亲二), A new morphological interpretation of the female reproductive organs in Ginkgo biloba L. with a phylogenetic consideration on gymnosperms. Acta Phytotax Sin (植物分类学报), 1993a, 31(3): 294-296
31. Fu DZ (傅德志)and Yang QE (杨亲二), A new morphological interpretation of the female reproductive organs in Ginkgo biloba L. with a phylogenetic consideration on gymnosperms (Cont.). Acta Phytotax Sin (植物分类学报), 1993b, 31(4): 309-317
32. Fu LC, Yu YF and Farjon A, Cupressaceae. In Wu ZY and Raven PH eds. Flora of China. Beijing: Science Press, St. Louis: Missouri Botanical Garden Press, 1999, 4: 62-77
33． Gadek PA, Alpers DL, Heslewood MM and Quinn CJ, Relationships within Cupressaceae sensu lato: a combined morphogical and molecular approach. Am J Bot, 2000, 87(8) : 1044-1057
34． Gadek PA & Quinn CJ, An analysis of relationships within the Cupressaceae sensu stricto based on rbcL sequences. Ann Missouri Bot Gard, 1993, 80: 581-588
35． Gadek PA & Quinn CJ, Biflavones of the subfamily Cupressoideae, Cupressaceae. Phytochemistry, 1985, 24: 267-272
36． Gadek PA & Quinn CJ, Pitting of transfusion tracheids in Cupressaceae. Austral J Bot, 1988, 36:81-92
37． Gifford EM and Foster AS, Morphology and evolution of vascular plants. New York, Freeman WH and Company. 1989．
38． Guedes M and Dupuy P, Morphology of the seed-scale-complex in Pinus pinaster subsp. atlanica: a teratologic approach. Phytomorphology, 1971, 21(1) : 17-31
39． Hart JA and Price RA, The genera of Cupressaceae (including Taxodiaceae) in the southeastern United Sates. J Arnold Arbor, 1990, 71: 275-322
40． Hart JA, A cladistic analysis of conifers: preliminary results. J Arnold Arb, 1987, 68: 269-307
41． Hasebe M, Evolution of reproductive organs in land plants. J Plant Res, 1999, 112(Dec): 463-474
42． Hayata B, The Sciadopityaceae represented by Sciadopitys verticillata Sieb et Zucc, an endemic species of Japan. Bot Mag (Tokyo), 1931, 45: 567-569
43． Hida M, The comparative study of Taxodiaceae from the standpoint of the development of the cone scale. Bot Mag (Tokyo), 1957, 70: 44-51
44． Hida M, The comparative study of Taxodiaceae from the standpoint of the development of the cone scale (supplement) the development of Metasequia's cone scale. Bot Mag (Tokyo), 1961, 74: 449-451
45． HuYS (胡玉熹) , Napp-Zinn K and Winne D, Comparative anatomy of seed-scale of female cones of Pinaceae. Bot Jahrb Syst, 1989, 111(1) : 63-85
46． Jain KK, Morphology of Female cone in Pinaceae. Phytomorphology, 1976, 26(2) : 189-200
47． Jain KK, Morphology of female cone in Podocarpus neriifolius. Phytomorphology, 1977,27:215-224
48． Keng H, Taxonomic position of Phyllocladus and the classification of conifers. Gardens' Bulletin Singapore, 1963, 20:127-130
49． Konar RN, Banerjee SK, The morphology and embryology of Cupressus funebris Endl. Phytomorphology, 1963a, 3: 321-338
50． Koner RN, Anatomical studies on Indian pines, with special references to Pinus
roxburghii Sar. Phytomorphology, 1963b, 13: 388-402．
51． Kurmann MH, Pollen morphology and ultrastructure in the Cupressaceae. Acta Bot Gallica, 1994, 141:141-147
52． Kusumi J, Tsumura Y, Yoshimaru H, and Tachida H, Phylogenetic relationships in Taxodiaceae and Cupressaceae sensu stricto based on matK gene, chlL gene, trnL-trnF IGS region and trnL intron sequences. Amer J Bot, 2000, 87(10) :1480-1488
53． Lemoine-Sebastian C, Etude comparative de la vascularisation et du complexe se minal chez les cupressacees. Phytomorphology, 1972, 22: 246-260
54． Lev-Yadun S and Liphschitz N, The ontogeny of gender of Cupressus sempervirens L., Bot Gaz, 1987, 148: 407-412
55． Lev-Yadun S, A develpmental variant of indentation in the scales of female coes of Cupressus sempervirens L. Bot J Linn Soc, 1996, 121: 263-269
56． Lev-Yadun S, Abnormal cones in Cupressus sempervirens. ZLISO, 1992, 13(2) : 391-394
57． Li HL, A reclassification of Libocedrus and Cupressaceae. J Arnold Arb, 1953, 34:17-35
58． Li SJ, The female reproductive organs of Chamaecyparis. Taiwania, 1972, 17(1) : 27-39
59． Longman KA, Dick J and Page CN, Cone induction with gibberellin for taxonomic studies in Cupressaceae and Taxodiaceae. Biol Pl, 1982, 24: 195-201
60． Marquard RD and Hanover JW, Sexual zonation in the crown of Picea glauca and the flowering response to exogenous GA4/7． Can J Forest Res, 1984, 14:27-30
61． Martin PC, A morphological comparison of Biota and Thuja. Proc Penn Acad Sci, 1950,24:65-112．
62． Master MT, Review of some point in the comparative morphology, anotomy and life history of the coniferae. Bot J Linn Soc, 1889, 27:226-332
63． Mc William JR, The role of the micropyle in the pollination of Pinus. Bot Gaz, 1958, 120:109-117
64． Meeuse ADJ, Fundamentals of Plant Morphology. New York: The Ronald Press Company. 1966．
65． Meeuse ADJ, From ovule to ovary: a contribution to the phylogeny of the megasporangium. Acta Biotheoretica, 1963, 16: 127-182
66． Meyen SV, Basic features of gymnosperm systematics and phylogeny as evidence by the fossil record. Bot Rev, 1984, 50:1
67． Miller CN, Implications of fossil conifers for the phylogenetic relationships of living families. Bot Rev, 1999, 65(3) : 239-277
68． Miller CN, The origin of morden conifer families. In Beck CB ed., Origin and Evolution of Gymnosperms. New York: Columbia University Press. 1988．
69． Morgan CS, Plant portraits: 368 Platycladus orientalis Cupressacese. Curtis's Bot Mag, 1999, 16(3) : 185-192
70． Mouradov A, Glassick T, Hamdorf B, Murphy L, Fowler B, Maria S and Teasdale RD, NEEDLY, a Pinus radiata ortholog of FLORICAULA/LEAFY genes, expressed in both reproductive and vegetative meristerms. Proc Natl Acad Sci USA. 1998, 94: 2415-2420
71． Napp-Zinn K and Hu YS (胡玉熹) , Anatomical studies on the bracts in pinaceous female cones III. Comparative study of (mostly Chinese) representatives of all genera. Bot Jahrb Syst, 1989, 110(4) : 461-477
72． Napp-Zinn K, Anatomie des Blattes. I. Blttananatomie der Gymnospermen. Gebruder Borntraeger, Berlin-Nikolassee. 1966．
73． Nelson T, Dengler N, Leaf vascular pattern formation. The Plant Cell, 1997, 9: 1121-1135
74． Ohsawa T, Anatomy and relationships of petrified seed cones of the Cupressaceae, Taxodiaceae and Sciadopityaceae. J Plant Res, 1994, 107: 503-512
75． Oladele FA, Inner surface sculpture patterns of cuticles in Cupressaceae. Can J Bot, 1983a, 61: 1222-1231
76． Oladele FA, Scanning electron microscope study of stomatal-complex configuration in Cupressaceae. Can J Bot, 1983b, 61: 1232-1240
77． Owens JN and Blake MD, Pollen morphology and development of the pollination mechanism in Tsuga heterophylla and T mertensiana. Can J Bot, 1984, 61:3041-3048
78． Owens J N and Molder M, Bud development in western hemlock Ⅱ, initiation and early development of pollen and seed cone. Can J Bot, 1974a, 52: 283-294
79． Owens J N and Molder M, Cone initiation and development before dormancy in yellow cedar (Chamaecyparis nootkatensis). Can J Bot, 1974b, 52: 2075-2084
80． Owens J N and Molder M, Sexual reproduction in western red cedar (Thuja plicata) Can J Bot, 1980, 58:1376-1393
81． Owens JN and Pharis RP, Initiation and development of western red cedar cones in response to gibberellin induction and under natural conditions. Can J Bot, 1971, 49: 1165-1175
82． Owens JN and Pharis RP, Initiation and ontogeny of the microsporangiate cone in Cupressus arizonica in response to gibberellin. Am J Bot, 1967, 54(10) : 1260-1272
83． Owens JN, Simpson SJ and Molder M, The pollination mechanism in yellow cypress (Chamaecypairis nootkatensis). Can J Forest Res, 1980, 10: 564-572
84． Owens JN, Simpson SJ and Molder M, Sexual reproduction of Pinus contora. I. Pollen development, the pollination mechanism and early ovule development. Can J Bot, 1981,59: 1828-1843
85． Owens JN, The relative importance of initiation and early development on cone production in Douglas fir. Can J Bot, 1969, 47: 1039-1049
86． Owens JN, Takaso T and Runions CJ, Pollination in conifers. Trends in Plant Science, 1998, 3(12) : 478-485
87． Page CN, Cupressaceae. In Kubitzki ed. The Families and Ggenera of Vascular Plants. Vol. 1 Pterdophytes and Gymnosperms. Springer-Verlag, 1990, pp302-316
88． Pant DD, The classificstion of gymnosperms plant. Paleobot, 1957, 6:65-70
89． Peirce AS, Systematic anatomy of the woods of the Cupressaceae. Trop Woods, 1937,49:5-21
90． Pharis RP and King RW, Gibberelins and reproductive development in seed plants. Ann Rev Pl Physiol, 1985, 36: 517-568
91． Pharis RP and Morf W, Experiments on the precocious flowering of western cedar and four species of Cupressus with gibberellins A_3, and A_4/A_7 mixture. Can J Bot, 1967,45:1519-1524
92． Pilger R, Gymnospermae. In: Engler A and Prantl K, Die naturlichen Pflanzenfamilien. Engelmann, Leipzing, 1926, 13: 121-407
93． Powell GR and Tosh KJ, The pollination mechanism and development after bud burst of cons of Lam laricina. Can J Bot, 1991, 69: 1468-1460
94． Price RA and Lowenstein JM, An immunological comparison of the Sciadopityaceae, Taxoidiaceae and Cupressaceae. Syst Bot, 1989, 14:141-149
95． Quinn CJ & Gadek PA, Sequence of xylem differentiation in leaves of Cupressaceae. Amer J Bot, 1988, 75(9) : 1344-1351
96． Radais M, Contribution I'etude de I'anatomic comparee du Fruit des Coniferers. Ann Sci Nat Bot, 1894, Ser. VII, 19: 165-368
97． Rutlelege R, Regan S, Nicalas O, Fobert P, Cote C, Bosnich W, Kauffeldt W, Sunohara G Seguin A and Stewart D, Characterization of an AGAMOUS homologue from the conifer black spruce (Picea mariana) that produces floral homeotic conversions when expressed in Arabidopsis. Plant J, 1998, 15: 625-634
98． Sahni B, On certain archaic features in the seed of Taxus baccata, with remarks on the antiquity of the Taxineae. Ann Bot, 1920, 34: 117-133
99． Satake Y, On the systematic importance of the vascular course in the cone scales of the Taxosiaceae (priliminary report). Bot Mag, 1934, XLVIII: 186-205
100． Sattler R and Rutishauser R, The fundamental relevance of morphology and morphogenesis to plant research. Ann Bot, 1997, 80: 571-582
101． Sattler R, Process morphology: structural dynamics in development and evolution. Can J Bot, 1992, 70: 708-714
102． Saxton WT, Contributions to the life history of Widdringtonia cupressoides. Bot Gaz, 1905, 50:31-48
103． Shi L and Wang FH, Development of the megasporangiate shoot of Taxaceae. Cathaya, 1989,1:83-91
104． Shindo S, Ito M, Ueda K, Kato M and Hasede M, Characterization of MADS genes in the gymnosperm Gnetum parvifolium and its implication on the evolution of reproductive organs in seed plants. 1999, Evol Dev, 1(3) : 1-11
105． Singh H and Oberoi YP, A contribution to the life history of Biota orientalis Endl. Phytomorphology, 1962, 12: 373-393
106． Singh H, Embryology of Gymnosperms. (Handbuch Pflanzenanatomie) Berlin: Borntraeger, 1978．
107． Sporne K R. The morphology of Gymnosperms. 2d ed. London: Hutchinson and Co. 1974．
108． Stefanoviac S, Jager M, Broutin J and Masselot M, Phylogenetic relationships of conifers inferred from partial 28S rRNA gene sequences. Am J Bot, 1998, 85: 688-697
109． Stockey RA, Some comment on the origin and evolution of conifers. Can J Bot, 1981,59: 1932-1940
110． Stoffberg E, Morphological and ontogenic studies on southern African Podocarpus, initiation of the seed scale complex and early development of integument, nucellus and epimatium. Bot J Lin Soc, 1991, 105: 21-35
111． Stutzel K and Rowekamp I, Female reproductive structures in Taxales. Flora, 1999, 194: 145-157
112． Sugihara Y, The embryogeny of Biota orientalis Endlicher and the secondary cleavage polyembryony in coniferales. J Bot, 1992, 67: 83-87
113． Takaso T and Owens JN, Ovulate cone morphology and pollination in Pseudotsuga and Cedrus. Int J Plant Sci 1995,156:630-639
114． Takaso T and Tomlinson PB, Aspect of cone and ovule ontogeny in Cryptomeria (Taxodiaceae) Am J Bot, 1989a, 76(5) : 692-705
115． Takaso T and Tomlinson PB, Cone and ovule development in Callitris (Cupressaceae-Callitroideae). Bot Gaz, 1989b, 150(4) : 378-390
116． Takaso T and Tomlinson PB, Cone and ovule ontogeny in Sciadopitys (Taxodiaceae-Coniferales). Am J Bot, 1991, 78: 417-428
117． Takaso T and Tomlinson PB, Cone and ovule ontogeny in Taxodium and Glyptostrobus (Taxodiaceae-Coniferales). Am. J. Bot., 1990, 77:1209-1211
118． Takaso T and Tomlinson PB, Seed cone and ovule ontogeny in Metasequoia, Sequoia and Sequiadendron (Taxodiaceae-Coniferales). Bot J Linn Soc, 1992, 109:15-37
119． Takaso T, A developmental study of the integument in gymnosperms (2) Pinns thunbergii Parl., Abies mariesii Mast. and A. veitchii Lindl. Jjap Bot, 1981, 56:73-89
120． Takhtajan AL, Phylogenetic principles of higher plants. Bot Rev, 1953, 19: 1-45
121． Tandre K, Svensson ME and Engstrom P, Conservation of gene structure and activity in the regulation of reproductive organ development of conifers and angiosperms. Plant J, 1998, 15; 615-623
122． Thomson RB, The structure of the cone in the coniferae. Bot Rev, 1940, 6: 73-84
123． Tomlinson PB and Takaso T, Hydrodynamics of pollen capture in conifers. In: Owens SJ and Rudall eds, Reproductive Biology. Royal Botanic Gardens Kew. 1998, pp. 265-275．
124． Tomlinson PB, Aspect of cone morphology and development in Podocarpaceae (Conoferales). Int J Plant Sci, 1992, 153(4) : 572-588
125． Tomlinson PB, Braggins JE and Rattenbury JA, Pollination drop in relation to cone morphology in Podocarpaceae: a novel reproductive mechanism. Am J Bot, 1991,78: 1289-1303
126． Tomlinson PB, Takaso T and Cameron EK, Cone development in Libocedrus (Cupressaceae)--phenological and morphological aspects. Am J Bot, 1993, 80(6) : 649-659
127． Tomlinson PB, Takaso T and Rattenury JA, Cone and ovule ontogeny in Phyllocladus (Podocarpaceae). Bot J Linn Soc, 1989, 99:209-221
128． Van Tieghem P, Anatomie comparee de la fleur femelle et du fruit des Cycadees, des Coniferes et des Gnetacees. Ann Sci Nat Bot, 1869, 5 serie,10: 269-304
129． Wilson EG, Thuja orientalis Linnaeus and Juniperus chinensis. J Arnold Abr, 1930, 11:98-100
130． Wilson EG, Thuja orientalis Linnaeus. J Arnold Abr, 1926, 7:70-74
131． Winter KU, Becker A, Munster T, Kim JT, Saedler H and Theissen G, MADS-box genes reveal that gnetophytes are more closely related to conifer than to flowering plants. Proc Natl Acad Sci USA, 1999, 96: 7342-7347
132． Worsdell WC, Observations on the vascular system of the female 'flowers' of Coniferae. Ann Bot, 1899,13: 527-548
133． Worsdell WC, The structure of female "flower" in Coniferae, an historical study. Ann Bot, 1900, 14; 39-82
134．Xing S P(邢树平)，Hu Y X(胡玉熹)，Chen Z K(陈祖铿)andL Lin J X(林金星)，The ovule development，pollination drop formation and pollination process in Taxus chinensis.Acta Bot Sinica(植物学报)，2000，42(2)：126-132
135．Xing S P(邢树平)，Zhang Q(张泉)，Hu Y X(胡玉熹)，Chen Z K(陈祖铿)and Lin J X(林金星)．The mechanism of pollination in Platycladus orientalis and Thuja occidentalis(Cupressacese)．Acta Bot Sinica(植物学报)，1999，41(2)：130-132
136．Yang Y(杨永)，Fu DZ(傅德志)，Review on the megastrobilus theories of conifers．Acta Phytotax Sin(植物分类学报)，2001，39(2)：169-191
137．白书农，植物开花研究：从生理学到遗传学。植物科学进展，1997，1：146-163
138．陈祖铿，王伏雄，福建柏的配子体发育。植物学报，1980，22(1)：6-22
139．陈祖铿，王伏雄，福建柏后期胚的发育。植物学报，1981a，23(4)：266-272
140．陈祖铿，王伏雄，福建柏早期胚的发育及其系统位置的商榷。植物分类学报，1981b，19(1)：24-28
141．冯旻，傅德志，梁汉兴，路安民，耧斗菜属花部形态的发生。植物学报，1995，37(10)：791-794
142．江泽平，王豁然，柏科分类和分布：亚科、族和属。植物分类学报，1997，35(3)：236-248
143．李林初，柏科的细胞分类学研究，云南植物研究，1998,20(2)：197-203
144．李林初，刘永强，王玉勤，刘钢，侧柏亚科三种植物的核型及其细胞分类学研究。云南植物研究，1996，18(4)：439-444
145．李林初，徐炳声，侧柏和福建柏染色体的核型的研究。云南植物研究，1984，6(4)：447-451
146．李正理译，K．伊稍著，种子植物解剖学。上海科学技术出版社，1982。
147．梁汉兴，三白草科花部发育及其形态学意义。植物分类学报，1994，32(5)：425-432
148．孙坤，陈家宽，陈之端，被子植物系统学中花发育研究的进展及对今后研究的思考。植物分类学报，1998，36(6)：558-568
149．塔赫他间(中国科学院植物研究所译)，高等植物，北京：科学出版社，1963
150．王伏雄，陈祖铿，裸子植物系统发育的几个问题。植物学通报，1983，1(1)：4-7
151．王伏雄，胡玉熹，植物学名词解释。科学出版社，1982
152．王海洋，胡玉熹，王伏雄，红豆杉科植物的雌性生殖器官的结构及其系统位置的商榷。植物学集刊，1994,7：42-47
153．邢树平，松杉类植物花粉发育和传粉生物学研究。2000，(博士论文)
154．姚璧君，胡玉熹，松柏类植物叶子的比较解剖观察。植物分类学报，1983，20(3)：275-294
155．郑万均，傅立国，中国植物志，第七卷。北京：科学出版社，1978
2． Aulenback KR and LePage BA, Taxodium wallissi sp. nov.: first occurrence of Taxodium from the upper cretaceous. Int J Plant Sci, 1998, 159(2) : 367-390
3． Axsmith BJ, Taylor TN and Taylor EL, A new fossil conifer from the Triassic of North America: implications for models of ovulate cone scale evolution. Int J Plant Sci, 1998, 159(2) : 358-366
4． Baird AM, The life history of Callitris. Phytomorphology, 1953, 3:258-284
5． Beck CB, Origin and Evolution of Gmnosperms. New York: Columbia University Press, 1988
6． Bessey CE, The morphology of the pine cone. Bot Gaz, 1902, 33:157-159
7． Biswas C and Johri BM, Gmnosperns. Berlin: Springer-Verlag. 1997．
8． Bouman F, The ovule. In: Johri BM ed. Embryology of Angiosperms. Berlin: Springer-Verlag, 1984, 123-157
9． Brunsfeld ST Soltis PS, Soltis DE, Gadek PA, Quinn CJ, Strenge DD and Banker TM, Phylogenetic relationships among the genera of Taxodiaceae and Cupressaceae: evidence from rbcl sequences. Sys Bot, 1994, 19(2) : 253-262
10． Buchholz JT, The embryogeny of conifers. In Duggar BM ed. Proc Fourth Int Congr Plant Sci (Ithaca, New York, 1926) , Menasha, Wisconsin: Geoge Banta Publishing Company, 1929, 1:359-392
11． Buchhholz JT, The classification of Coniferales. Transaction of the Illinois State Academy of Science, 1934,25: 112-113
12． Chamberlain CJ, Gymnosperm Structure and Evolution. Chicago: The University of Chicago Press. 1935
13． Chaw SM, Zharkikh A, Sung HM et al, Molecular phylogeny of extant gymnosperm and seed plant evolution: analysis of nuclear 18S rRNA sequences. Mol Biol Evol, 1997, 14(1) : 56-68
14． Coen E and Meyerowitz EM, The war of the whorls: genetic interactions controlling flower development. Nature, 1991, 353: 31-37
15． Colombo L, Franken J, Koetje E et.al, The Petunia MADS-box gene FBP11 determines ovule identity. Plant Cell, 1995, 7: 1859-1868
16． Coulter PR and Chamberlain CJ, Morphology of Gymnosperms. Chicago: The University of Chicago Press. 1917．
17． Dalimore W and Jackson AB, A Handbook of Coniferae and Ginkgoaceae, 4th ed.
18. De Laubenfels DJ, Commentary: the organization of female fertile structure in conifers. Int J Plant Sci, 1992, 153 (4): ⅶ-ⅷ
19. De Laubenfels DJ, The relationships of Fitzroya cupressoides (Molina) Johnston and Diselma archeri J. D. Hooker based on morphological considerations. Phytomorphology, 1965, 15 (4): 414-419
20. Del Fueyo GM, Cone and ovule development in the Podocarpus species from Argentina. Phytomorphology. 1999, 49 (1): 49-60
21. Dogra PD, Embryogeny of gymnosperm and taxonomy ---- an assessment. In Nair PKK ed., Glimpses in Plant Research ---- Modern Trends in Plant Taxonomy, 1980, 5: 114-128
22. Eckenwalder JE, Comments on" A new classification of the conifers". Taxon, 1976a, 25: 337-338
23. Eckenwalder JE, Re-evaluation of Cupressaceae and Taxodiaceae: a proposed merger. Madroo, 1976b, 23 (5): 237-300
24. Endress PK, Diversity and evolutionary biology of tropical flowers. Cambridge University Press. 1994.
25. Farjon A, World Checklist and Bibliography of Conifers. The Royal Botanic Gardens, Kew. 1998
26. Florin R, Evolution in cordaites and confers. Acta Horti Bergiani, 1951, 15: 285-388
27. Florin R, The female reproductive organs of conifers and taxads. Bio Rev, 1954, 29: 367-389
28. Foster AS, Gifford EM, Comparative Morphology of Vascular Plants. New York: WH Freeman and Company, 1974.
29. Fu DZ (傅德志), Nageiaceae ---- a new gymnosperm family. Acta Phytotax Sin (植物分类学报), 1992, 30(6): 515-528
30. Fu DZ (傅德志)and Yang QE (杨亲二), A new morphological interpretation of the female reproductive organs in Ginkgo biloba L. with a phylogenetic consideration on gymnosperms. Acta Phytotax Sin (植物分类学报), 1993a, 31(3): 294-296
31. Fu DZ (傅德志)and Yang QE (杨亲二), A new morphological interpretation of the female reproductive organs in Ginkgo biloba L. with a phylogenetic consideration on gymnosperms (Cont.). Acta Phytotax Sin (植物分类学报), 1993b, 31(4): 309-317
32. Fu LC, Yu YF and Farjon A, Cupressaceae. In Wu ZY and Raven PH eds. Flora of China. Beijing: Science Press, St. Louis: Missouri Botanical Garden Press, 1999, 4: 62-77
33． Gadek PA, Alpers DL, Heslewood MM and Quinn CJ, Relationships within Cupressaceae sensu lato: a combined morphogical and molecular approach. Am J Bot, 2000, 87(8) : 1044-1057
34． Gadek PA & Quinn CJ, An analysis of relationships within the Cupressaceae sensu stricto based on rbcL sequences. Ann Missouri Bot Gard, 1993, 80: 581-588
35． Gadek PA & Quinn CJ, Biflavones of the subfamily Cupressoideae, Cupressaceae. Phytochemistry, 1985, 24: 267-272
36． Gadek PA & Quinn CJ, Pitting of transfusion tracheids in Cupressaceae. Austral J Bot, 1988, 36:81-92
37． Gifford EM and Foster AS, Morphology and evolution of vascular plants. New York, Freeman WH and Company. 1989．
38． Guedes M and Dupuy P, Morphology of the seed-scale-complex in Pinus pinaster subsp. atlanica: a teratologic approach. Phytomorphology, 1971, 21(1) : 17-31
39． Hart JA and Price RA, The genera of Cupressaceae (including Taxodiaceae) in the southeastern United Sates. J Arnold Arbor, 1990, 71: 275-322
40． Hart JA, A cladistic analysis of conifers: preliminary results. J Arnold Arb, 1987, 68: 269-307
41． Hasebe M, Evolution of reproductive organs in land plants. J Plant Res, 1999, 112(Dec): 463-474
42． Hayata B, The Sciadopityaceae represented by Sciadopitys verticillata Sieb et Zucc, an endemic species of Japan. Bot Mag (Tokyo), 1931, 45: 567-569
43． Hida M, The comparative study of Taxodiaceae from the standpoint of the development of the cone scale. Bot Mag (Tokyo), 1957, 70: 44-51
44． Hida M, The comparative study of Taxodiaceae from the standpoint of the development of the cone scale (supplement) the development of Metasequia's cone scale. Bot Mag (Tokyo), 1961, 74: 449-451
45． HuYS (胡玉熹) , Napp-Zinn K and Winne D, Comparative anatomy of seed-scale of female cones of Pinaceae. Bot Jahrb Syst, 1989, 111(1) : 63-85
46． Jain KK, Morphology of Female cone in Pinaceae. Phytomorphology, 1976, 26(2) : 189-200
47． Jain KK, Morphology of female cone in Podocarpus neriifolius. Phytomorphology, 1977,27:215-224
48． Keng H, Taxonomic position of Phyllocladus and the classification of conifers. Gardens' Bulletin Singapore, 1963, 20:127-130
49． Konar RN, Banerjee SK, The morphology and embryology of Cupressus funebris Endl. Phytomorphology, 1963a, 3: 321-338
50． Koner RN, Anatomical studies on Indian pines, with special references to Pinus
roxburghii Sar. Phytomorphology, 1963b, 13: 388-402．
51． Kurmann MH, Pollen morphology and ultrastructure in the Cupressaceae. Acta Bot Gallica, 1994, 141:141-147
52． Kusumi J, Tsumura Y, Yoshimaru H, and Tachida H, Phylogenetic relationships in Taxodiaceae and Cupressaceae sensu stricto based on matK gene, chlL gene, trnL-trnF IGS region and trnL intron sequences. Amer J Bot, 2000, 87(10) :1480-1488
53． Lemoine-Sebastian C, Etude comparative de la vascularisation et du complexe se minal chez les cupressacees. Phytomorphology, 1972, 22: 246-260
54． Lev-Yadun S and Liphschitz N, The ontogeny of gender of Cupressus sempervirens L., Bot Gaz, 1987, 148: 407-412
55． Lev-Yadun S, A develpmental variant of indentation in the scales of female coes of Cupressus sempervirens L. Bot J Linn Soc, 1996, 121: 263-269
56． Lev-Yadun S, Abnormal cones in Cupressus sempervirens. ZLISO, 1992, 13(2) : 391-394
57． Li HL, A reclassification of Libocedrus and Cupressaceae. J Arnold Arb, 1953, 34:17-35
58． Li SJ, The female reproductive organs of Chamaecyparis. Taiwania, 1972, 17(1) : 27-39
59． Longman KA, Dick J and Page CN, Cone induction with gibberellin for taxonomic studies in Cupressaceae and Taxodiaceae. Biol Pl, 1982, 24: 195-201
60． Marquard RD and Hanover JW, Sexual zonation in the crown of Picea glauca and the flowering response to exogenous GA4/7． Can J Forest Res, 1984, 14:27-30
61． Martin PC, A morphological comparison of Biota and Thuja. Proc Penn Acad Sci, 1950,24:65-112．
62． Master MT, Review of some point in the comparative morphology, anotomy and life history of the coniferae. Bot J Linn Soc, 1889, 27:226-332
63． Mc William JR, The role of the micropyle in the pollination of Pinus. Bot Gaz, 1958, 120:109-117
64． Meeuse ADJ, Fundamentals of Plant Morphology. New York: The Ronald Press Company. 1966．
65． Meeuse ADJ, From ovule to ovary: a contribution to the phylogeny of the megasporangium. Acta Biotheoretica, 1963, 16: 127-182
66． Meyen SV, Basic features of gymnosperm systematics and phylogeny as evidence by the fossil record. Bot Rev, 1984, 50:1
67． Miller CN, Implications of fossil conifers for the phylogenetic relationships of living families. Bot Rev, 1999, 65(3) : 239-277
68． Miller CN, The origin of morden conifer families. In Beck CB ed., Origin and Evolution of Gymnosperms. New York: Columbia University Press. 1988．
69． Morgan CS, Plant portraits: 368 Platycladus orientalis Cupressacese. Curtis's Bot Mag, 1999, 16(3) : 185-192
70． Mouradov A, Glassick T, Hamdorf B, Murphy L, Fowler B, Maria S and Teasdale RD, NEEDLY, a Pinus radiata ortholog of FLORICAULA/LEAFY genes, expressed in both reproductive and vegetative meristerms. Proc Natl Acad Sci USA. 1998, 94: 2415-2420
71． Napp-Zinn K and Hu YS (胡玉熹) , Anatomical studies on the bracts in pinaceous female cones III. Comparative study of (mostly Chinese) representatives of all genera. Bot Jahrb Syst, 1989, 110(4) : 461-477
72． Napp-Zinn K, Anatomie des Blattes. I. Blttananatomie der Gymnospermen. Gebruder Borntraeger, Berlin-Nikolassee. 1966．
73． Nelson T, Dengler N, Leaf vascular pattern formation. The Plant Cell, 1997, 9: 1121-1135
74． Ohsawa T, Anatomy and relationships of petrified seed cones of the Cupressaceae, Taxodiaceae and Sciadopityaceae. J Plant Res, 1994, 107: 503-512
75． Oladele FA, Inner surface sculpture patterns of cuticles in Cupressaceae. Can J Bot, 1983a, 61: 1222-1231
76． Oladele FA, Scanning electron microscope study of stomatal-complex configuration in Cupressaceae. Can J Bot, 1983b, 61: 1232-1240
77． Owens JN and Blake MD, Pollen morphology and development of the pollination mechanism in Tsuga heterophylla and T mertensiana. Can J Bot, 1984, 61:3041-3048
78． Owens J N and Molder M, Bud development in western hemlock Ⅱ, initiation and early development of pollen and seed cone. Can J Bot, 1974a, 52: 283-294
79． Owens J N and Molder M, Cone initiation and development before dormancy in yellow cedar (Chamaecyparis nootkatensis). Can J Bot, 1974b, 52: 2075-2084
80． Owens J N and Molder M, Sexual reproduction in western red cedar (Thuja plicata) Can J Bot, 1980, 58:1376-1393
81． Owens JN and Pharis RP, Initiation and development of western red cedar cones in response to gibberellin induction and under natural conditions. Can J Bot, 1971, 49: 1165-1175
82． Owens JN and Pharis RP, Initiation and ontogeny of the microsporangiate cone in Cupressus arizonica in response to gibberellin. Am J Bot, 1967, 54(10) : 1260-1272
83． Owens JN, Simpson SJ and Molder M, The pollination mechanism in yellow cypress (Chamaecypairis nootkatensis). Can J Forest Res, 1980, 10: 564-572
84． Owens JN, Simpson SJ and Molder M, Sexual reproduction of Pinus contora. I. Pollen development, the pollination mechanism and early ovule development. Can J Bot, 1981,59: 1828-1843
85． Owens JN, The relative importance of initiation and early development on cone production in Douglas fir. Can J Bot, 1969, 47: 1039-1049
86． Owens JN, Takaso T and Runions CJ, Pollination in conifers. Trends in Plant Science, 1998, 3(12) : 478-485
87． Page CN, Cupressaceae. In Kubitzki ed. The Families and Ggenera of Vascular Plants. Vol. 1 Pterdophytes and Gymnosperms. Springer-Verlag, 1990, pp302-316
88． Pant DD, The classificstion of gymnosperms plant. Paleobot, 1957, 6:65-70
89． Peirce AS, Systematic anatomy of the woods of the Cupressaceae. Trop Woods, 1937,49:5-21
90． Pharis RP and King RW, Gibberelins and reproductive development in seed plants. Ann Rev Pl Physiol, 1985, 36: 517-568
91． Pharis RP and Morf W, Experiments on the precocious flowering of western cedar and four species of Cupressus with gibberellins A_3, and A_4/A_7 mixture. Can J Bot, 1967,45:1519-1524
92． Pilger R, Gymnospermae. In: Engler A and Prantl K, Die naturlichen Pflanzenfamilien. Engelmann, Leipzing, 1926, 13: 121-407
93． Powell GR and Tosh KJ, The pollination mechanism and development after bud burst of cons of Lam laricina. Can J Bot, 1991, 69: 1468-1460
94． Price RA and Lowenstein JM, An immunological comparison of the Sciadopityaceae, Taxoidiaceae and Cupressaceae. Syst Bot, 1989, 14:141-149
95． Quinn CJ & Gadek PA, Sequence of xylem differentiation in leaves of Cupressaceae. Amer J Bot, 1988, 75(9) : 1344-1351
96． Radais M, Contribution I'etude de I'anatomic comparee du Fruit des Coniferers. Ann Sci Nat Bot, 1894, Ser. VII, 19: 165-368
97． Rutlelege R, Regan S, Nicalas O, Fobert P, Cote C, Bosnich W, Kauffeldt W, Sunohara G Seguin A and Stewart D, Characterization of an AGAMOUS homologue from the conifer black spruce (Picea mariana) that produces floral homeotic conversions when expressed in Arabidopsis. Plant J, 1998, 15: 625-634
98． Sahni B, On certain archaic features in the seed of Taxus baccata, with remarks on the antiquity of the Taxineae. Ann Bot, 1920, 34: 117-133
99． Satake Y, On the systematic importance of the vascular course in the cone scales of the Taxosiaceae (priliminary report). Bot Mag, 1934, XLVIII: 186-205
100． Sattler R and Rutishauser R, The fundamental relevance of morphology and morphogenesis to plant research. Ann Bot, 1997, 80: 571-582
101． Sattler R, Process morphology: structural dynamics in development and evolution. Can J Bot, 1992, 70: 708-714
102． Saxton WT, Contributions to the life history of Widdringtonia cupressoides. Bot Gaz, 1905, 50:31-48
103． Shi L and Wang FH, Development of the megasporangiate shoot of Taxaceae. Cathaya, 1989,1:83-91
104． Shindo S, Ito M, Ueda K, Kato M and Hasede M, Characterization of MADS genes in the gymnosperm Gnetum parvifolium and its implication on the evolution of reproductive organs in seed plants. 1999, Evol Dev, 1(3) : 1-11
105． Singh H and Oberoi YP, A contribution to the life history of Biota orientalis Endl. Phytomorphology, 1962, 12: 373-393
106． Singh H, Embryology of Gymnosperms. (Handbuch Pflanzenanatomie) Berlin: Borntraeger, 1978．
107． Sporne K R. The morphology of Gymnosperms. 2d ed. London: Hutchinson and Co. 1974．
108． Stefanoviac S, Jager M, Broutin J and Masselot M, Phylogenetic relationships of conifers inferred from partial 28S rRNA gene sequences. Am J Bot, 1998, 85: 688-697
109． Stockey RA, Some comment on the origin and evolution of conifers. Can J Bot, 1981,59: 1932-1940
110． Stoffberg E, Morphological and ontogenic studies on southern African Podocarpus, initiation of the seed scale complex and early development of integument, nucellus and epimatium. Bot J Lin Soc, 1991, 105: 21-35
111． Stutzel K and Rowekamp I, Female reproductive structures in Taxales. Flora, 1999, 194: 145-157
112． Sugihara Y, The embryogeny of Biota orientalis Endlicher and the secondary cleavage polyembryony in coniferales. J Bot, 1992, 67: 83-87
113． Takaso T and Owens JN, Ovulate cone morphology and pollination in Pseudotsuga and Cedrus. Int J Plant Sci 1995,156:630-639
114． Takaso T and Tomlinson PB, Aspect of cone and ovule ontogeny in Cryptomeria (Taxodiaceae) Am J Bot, 1989a, 76(5) : 692-705
115． Takaso T and Tomlinson PB, Cone and ovule development in Callitris (Cupressaceae-Callitroideae). Bot Gaz, 1989b, 150(4) : 378-390
116． Takaso T and Tomlinson PB, Cone and ovule ontogeny in Sciadopitys (Taxodiaceae-Coniferales). Am J Bot, 1991, 78: 417-428
117． Takaso T and Tomlinson PB, Cone and ovule ontogeny in Taxodium and Glyptostrobus (Taxodiaceae-Coniferales). Am. J. Bot., 1990, 77:1209-1211
118． Takaso T and Tomlinson PB, Seed cone and ovule ontogeny in Metasequoia, Sequoia and Sequiadendron (Taxodiaceae-Coniferales). Bot J Linn Soc, 1992, 109:15-37
119． Takaso T, A developmental study of the integument in gymnosperms (2) Pinns thunbergii Parl., Abies mariesii Mast. and A. veitchii Lindl. Jjap Bot, 1981, 56:73-89
120． Takhtajan AL, Phylogenetic principles of higher plants. Bot Rev, 1953, 19: 1-45
121． Tandre K, Svensson ME and Engstrom P, Conservation of gene structure and activity in the regulation of reproductive organ development of conifers and angiosperms. Plant J, 1998, 15; 615-623
122． Thomson RB, The structure of the cone in the coniferae. Bot Rev, 1940, 6: 73-84
123． Tomlinson PB and Takaso T, Hydrodynamics of pollen capture in conifers. In: Owens SJ and Rudall eds, Reproductive Biology. Royal Botanic Gardens Kew. 1998, pp. 265-275．
124． Tomlinson PB, Aspect of cone morphology and development in Podocarpaceae (Conoferales). Int J Plant Sci, 1992, 153(4) : 572-588
125． Tomlinson PB, Braggins JE and Rattenbury JA, Pollination drop in relation to cone morphology in Podocarpaceae: a novel reproductive mechanism. Am J Bot, 1991,78: 1289-1303
126． Tomlinson PB, Takaso T and Cameron EK, Cone development in Libocedrus (Cupressaceae)--phenological and morphological aspects. Am J Bot, 1993, 80(6) : 649-659
127． Tomlinson PB, Takaso T and Rattenury JA, Cone and ovule ontogeny in Phyllocladus (Podocarpaceae). Bot J Linn Soc, 1989, 99:209-221
128． Van Tieghem P, Anatomie comparee de la fleur femelle et du fruit des Cycadees, des Coniferes et des Gnetacees. Ann Sci Nat Bot, 1869, 5 serie,10: 269-304
129． Wilson EG, Thuja orientalis Linnaeus and Juniperus chinensis. J Arnold Abr, 1930, 11:98-100
130． Wilson EG, Thuja orientalis Linnaeus. J Arnold Abr, 1926, 7:70-74
131． Winter KU, Becker A, Munster T, Kim JT, Saedler H and Theissen G, MADS-box genes reveal that gnetophytes are more closely related to conifer than to flowering plants. Proc Natl Acad Sci USA, 1999, 96: 7342-7347
132． Worsdell WC, Observations on the vascular system of the female 'flowers' of Coniferae. Ann Bot, 1899,13: 527-548
133． Worsdell WC, The structure of female "flower" in Coniferae, an historical study. Ann Bot, 1900, 14; 39-82
134．Xing S P(邢树平)，Hu Y X(胡玉熹)，Chen Z K(陈祖铿)andL Lin J X(林金星)，The ovule development，pollination drop formation and pollination process in Taxus chinensis.Acta Bot Sinica(植物学报)，2000，42(2)：126-132
135．Xing S P(邢树平)，Zhang Q(张泉)，Hu Y X(胡玉熹)，Chen Z K(陈祖铿)and Lin J X(林金星)．The mechanism of pollination in Platycladus orientalis and Thuja occidentalis(Cupressacese)．Acta Bot Sinica(植物学报)，1999，41(2)：130-132
136．Yang Y(杨永)，Fu DZ(傅德志)，Review on the megastrobilus theories of conifers．Acta Phytotax Sin(植物分类学报)，2001，39(2)：169-191
137．白书农，植物开花研究：从生理学到遗传学。植物科学进展，1997，1：146-163
138．陈祖铿，王伏雄，福建柏的配子体发育。植物学报，1980，22(1)：6-22
139．陈祖铿，王伏雄，福建柏后期胚的发育。植物学报，1981a，23(4)：266-272
140．陈祖铿，王伏雄，福建柏早期胚的发育及其系统位置的商榷。植物分类学报，1981b，19(1)：24-28
141．冯旻，傅德志，梁汉兴，路安民，耧斗菜属花部形态的发生。植物学报，1995，37(10)：791-794
142．江泽平，王豁然，柏科分类和分布：亚科、族和属。植物分类学报，1997，35(3)：236-248
143．李林初，柏科的细胞分类学研究，云南植物研究，1998,20(2)：197-203
144．李林初，刘永强，王玉勤，刘钢，侧柏亚科三种植物的核型及其细胞分类学研究。云南植物研究，1996，18(4)：439-444
145．李林初，徐炳声，侧柏和福建柏染色体的核型的研究。云南植物研究，1984，6(4)：447-451
146．李正理译，K．伊稍著，种子植物解剖学。上海科学技术出版社，1982。
147．梁汉兴，三白草科花部发育及其形态学意义。植物分类学报，1994，32(5)：425-432
148．孙坤，陈家宽，陈之端，被子植物系统学中花发育研究的进展及对今后研究的思考。植物分类学报，1998，36(6)：558-568
149．塔赫他间(中国科学院植物研究所译)，高等植物，北京：科学出版社，1963
150．王伏雄，陈祖铿，裸子植物系统发育的几个问题。植物学通报，1983，1(1)：4-7
151．王伏雄，胡玉熹，植物学名词解释。科学出版社，1982
152．王海洋，胡玉熹，王伏雄，红豆杉科植物的雌性生殖器官的结构及其系统位置的商榷。植物学集刊，1994,7：42-47
153．邢树平，松杉类植物花粉发育和传粉生物学研究。2000，(博士论文)
154．姚璧君，胡玉熹，松柏类植物叶子的比较解剖观察。植物分类学报，1983，20(3)：275-294
155．郑万均，傅立国，中国植物志，第七卷。北京：科学出版社，1978